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UNITED  STATES  DEPARTMENT  OF  THE  INTERIOR,  Douglas  McKay,  Secretary    J2 
FISH  AND  WILDLIFE  SERVICE,  John  L.  Farley,  Director 


FISHES  OF  THE  GULF  OF  MAINE 


BY  HENRY  B.  BIGELOW  AND  WILLIAM  G.  SCHROEDER 


First  Revision 


FISHERY  BULLETIN  74 


FISHERY  BULLETIN  OF  THE  FISH  AND  WILDLIFE  SERVICE 

VOLUME  53 

[Contribution  No.  S92,  Woods  Hole  Oceanographic  Institution} 


UNITED  STATES  GOVERNMENT  PRINTING  OFFICE  •  Washington  :  1953 


NOTICE 

Bigelow  and  Schroeder's  "Fishes  of  the  Gulf  of  Maine"  was  printed  in 
1953  and  went  on  sale  at  the  U.S.  Government  Printing  Office  on  February  12, 
1954.  This  was  a  revision  of  an  earlier  work  of  the  same  name  by  Bigelow  and 
W.  W.  Welsh  (1925);  3,493  copies  of  the  revision  were  printed.  Of  these, 
2,000  copies  were  distributed  by  the  Fish  and  Wildlife  Service,  the  remainder 
by  the  Government  Printing  Office.  The  Service  supply  was  exhausted  in  Janu- 
ary 1961  and  that  of  the  Printing  Office  in  March  1961. 

A  photo-offset  reprinting  was  issued  in  1964  jointly  by  the  Woods  Hole 
Oceanographic  Institution  and  the  Museum  of  Comparative  Zoology,  Harvard 
University,  organizations  with  which  the  authors  have  been  associated  for  many 
years.  Since  then,  additional  reprintings  have  followed.  The  contents  of  the  book 
are  precisely  the  same  as  published  in  1953  save  for  the  addition  of  this  note 
and  the  deletion  of  two  lines  at  the  foot  of  the  title  page  saying  "For  sale  by  the 
Superintendent  of  Documents,  U.S.  Government  Printing  Office,  Washington  25, 
D.C.  Price  $4.25  (Buckram)." 

This  reprint  may  be  obtained  from  the  Museum  of  Comparative  Zoology, 
Harvard  University,  Cambridge,  Massachusetts  02138. 


CONTENTS 

Page 

Introduction 1 

Area  covered 1 

Scope  of  the  work 1 

Sources  of  information 2 

Use  of  the  keys 4 

Key  to  Gulf  of  Maine  fishes 5 

The  Cyclostomes.     Class  Agnatha 9 

Hagfishes  and  lampreys.     Families  Myxinidae  and  Petromyzonidae 9 

Hag  Myxine  glutinosa  Linnaeus 10 

Sea  lamprey  Petromyzon  marinus  Linnaeus 12 

Cartilaginous  fishes.     Class  Chondrichthyes 15 

Sharks,  torpedoes,  skates  and  rays.     Subclass  Elasmobranchii 15 

Sharks.     Order  Selachii 15 

Sand  sharks.     Family  Carchariidae 18 

Sand  shark  Carcharias  taurus  Rafinesque 18 

Mackerel  sharks.     Family  Isuridae 20 

Mackerel  shark  Lamna  nasus  (Bonnaterre) 20 

Sharp-nosed  mackerel  shark,  Mako  I  sums  oxyrinchus  Rafinesque 23 

Maneater,  White  shark  Carcharodon  carcharias  (Linnaeus) 25 

Basking  sharks.     Family  Cetorhinidae 28 

Basking  shark  Celorhinus  maximus  (Gunnerus) 28 

Thresher  sharks.     Family  Alopiidae 32 

Thresher  Alopias  vulpinus  (Bonnaterre) 32 

Cat  sharks.     Family  Scyliorhinidae 34 

Chain  dogfish  Scyliorhinus  retifer  (Garman) 34 

Smooth  dogfishes.     Family  Triakidae 34 

Smooth  dogfish  Mustelus  canis  (Mitchill) 34 

Requiem  sharks.     Family  Carcharhinidae 36 

Tiger  shark  Galeocerdo  cuvier  (LeSueur) 37 

Blue  shark  Prionace  glauca  (Linnaeus) 38 

Sharp-nosed  shark  Scoliodon  terrae-novae  (Richardson) 40 

Dusky  shark  (Carcharhinus  obscurus  (LeSueur) 41 

Brown  shark  Carcharhinus  milberti  (Muller  and  Henle) 43 

Hammerhead  sharks.     Family  Sphyrnidae 44 

Bonnet  shark,  Shovelhead  Sphyrna  tiburo  (Linnaeus) 44 

Hammerhead  Sphyrna  zygaena  (Linnaeus) 45 

Spiny  dogfishes.     Family  Squalidae 47 

Spiny  dogfish  Squalus  acanthias  Linnaeus 47 

Black  dogfish  Centroscyllium  fabricii  (Reinhardt) 51 

Portuguese  shark  Cenlroscymnus  coelolepis  Bocage  and  Brito  Capello 52 

Gurry  sharks.     Family  Dalatiidae 53 

Greenland  shark  Somniosus  microcephalus  (Bloch  and  Schneider) 53 

Dalalias  licha  (Bonnaterre) 55 

Bramble  sharks.     Family  Echinorhinidae 56 

Bramble  shark  Echinorhinus  brucus  (Bonnaterre) 56 

Torpedoes,  skates  and  rays.     Order  Batoidei 57 

Torpedo  or  electric  rays.     Family  Torpedinidae 58 

Torpedo  Torpedo  nobiliana  Bonaparte 58 

Skates.     Family  Rajidae 60 

Barn-door  skate  Raja  laevis  Mitchill 61 

Big  skate  Raja  ocellata  Mitchill 63 

Brier  skate  Raja  eglanteria  Bosc 65 

Leopard  skate  Raja  garmani  Whitley 66 

Little  skate  Raja  erinacea  Mitchill 67 

Smooth- tailed  skate  Raja  senla  Garman 70 

Thorny  skate  Raja  radiata  Donovan 72 


m 


jy  CONTENTS 

Cartilaginous  fishes.     Class  Chondrichthyes — Continued 

Sharks,  torpedoes,  skates  and  rays.     Subclass  Elasmobranchii — Continued 

Torpedoes,  skates  and,  rays.     Order  Batoidei — Continued  Page 

Whip-tailed  sting  rays.     Family  Dasyatidae 74 

Sting  ray  Dasyatis  centroura  (Mitchill) 74 

Cow-nosed  rays.     Family  Rhinopteridae 76 

Cow-nosed  ray  Rhinoptera  bonasus  ( Mitchill) 76 

Devil  rays.     Family  Mobulidae 77 

Devil  ray  Manta  birostris  (Donndorff) 77 

Chimaeroids.     Subclass  Holocephali 79 

Chimaeras.     Order  Chimaerae 79 

Family  Chimaeridae 79 

Chimaera  Hydrolagus  affinis  (Brito  Capello) 79 

Bony  fishes.     Class  Osteichthyes 80 

Sturgeons.     Family  Acipenseridae 80 

Sea  sturgeon  Acipenser  sturio  Linnaeus 81 

Short-nosed  sturgeon  Acipenser  brevirostrum  LeSueur 84 

Herring  and  Tarpon  tribes.     Families  Clupeidae,  Dussumieriidae,  and  Elopidae.  85 

Ten-pounder  Elops  saurus  Linnaeus 86 

Tarpon  Tarpon  atlanticus  (Cuvier  and  Valenciennes) 87 

Round  herring  Etrumeus  sadina  (Mitchill) 87 

Herring  Clupea  harengus  Linnaeus 88 

Hickory  shad  Pomolobus  mediocris  (Mitchill) 100 

Alewife  Pomolobus  pseudoharengus  (Wilson) 101 

Blueback  Pomolobus  aestivalis  (Mitchill) 106 

Shad  Alosa  sapidissima  (Wilson) 108 

Thread  herring  Opislhonema  oglinum  (LeSueur) 112 

Menhaden  Brevoortia  tyrannus  (Latrobe) 113 

Anchovies.     Family  Engraulidae 118 

Anchovy  Anchoa  mitchilli  (Cuvier  and  Valenciennes) 118 

Striped  anchovy  Anchoa  hepsetus  (Linnaeus) 119 

Salmons.     Family  Salmonidae H» 

Brook  trout  Salvelinus  fontinalis  (Mitchill) 120 

Salmon  Salmo  salar  Linnaeus 121 

Humpback  salmon  Oncorhynchus  gorbuscha  (Walbaum) 131 

Silver  salmon  Oncorhynchus  kisulch  (Walbaum) 133 

Smelts.     Family  Osmeridae 133 

Capelin  Mallotus  villosus  (Mtiller) 134 

Smelt  Osmerus  mordax  (Mitchill) 135 

Argentines.     Family  Argentinidae 139 

Argentine  Argentina  silus  Ascanius 139 

Luminescent  fishes ™* 

Lanternfishes.     Family  Myctophidae 141 

Headlight-fish  Diaphus  effulgens  (Goode  and  Bean) 142 

Lanternfish  Myctophum  affine  (Liitken) 143 

Pearlsides.     Family  Maurolicidae 143 

Pearlsides  Maurolicus  pennanti  (Walbaum) 144 

Viper  Fishes.     Family  Chauliodontidae 145 

Viperfish  Chauliodus  sloani  Bloch  and  Schneider 145 

Stomiatids.     Families  Gonostomidae  and  Stomiatidae 146 

Cyclothone  Cyclothone  signata  Garman 146 

Stomias  Stomias  ferox  Rheinhardt 147 

Stomioides  nicholsi  Parr 147 

Trigonolampa  miriceps  Regan  and  Trewavas 148 

Hatchet  fishes.     Family  Sternoptychidae 149 

Silver  hatchet  fish  Argyropelecus  aculealus  Cuvier  and  Valenciennes 149 

Eels.     Families  Anguillidae,  Congridae,  Simenchelyidae,  Synaphobranchidae, 

Nemichthyidae,  and  Ophichthyidae 150 

Eel  Anguilla  rostrata  (LeSueur) 151 

American  conger  Conger  oceanica  ( Mitchill) 154 


CONTENTS 

Bony  fishes.     Class  Osteichthyes — Continued  Paee 

Eels — Continued 

Slime  eel  Simenchelys  parasiticus  Gill 157 

Long-nosed  eel  Synaphobranchus  pinnatus  (Gronow) 158 

Snake  eel  Omochelys  cruentifer  (Goode  and  Bean) 159 

Snipe  eel  Nemichthys  scolopaceus  Richardson 159 

Lancetfishes.     Family  Alepisauridae 160 

Lancetfish  Alepisaurus  ferox  Lowe 161 

Mummichogs  or  killifishes.     Family  Poeciliidae 162 

Common  mummichog  Fundulus  heteroclitus  (Linnaeus) 162 

Striped  mummichog  Fundulus  majalis  (Walbaum) 164 

Sheepshead  minnow  Cyprinodon  variegatus  Lacepede 165 

Billfishes.     Family  Belonidae 167 

Silver  gar  Tylosurus  marinus  (Walbaum) 167 

Garfish  Ablennes  hians  (Cuvier  and  Valenciennes) 168 

Halfbeaks.     Family  Hemiramphidae 169 

Halfbeak  Hyporhamphus  unifasciatus  (Ranzani) 169 

Needlefishes.     Family  Scomberesocidae 170 

Needlefish  Scomberesox  saurus  (Walbaum) 170 

Flying  fishes.     Family  Exocoetidae 171 

Flying  fish.     Cypselurus  heterurus  (Rafinesque) 172 

Silver  hake  and  Cod  families.     Families  Merlucciidae  and  Gadidae 173 

Silver  hake  Merluccius  bilinearis  (Mitchill) 173 

Cod  Gadus  callarias  Linnaeus 182 

Tomcod  Microgadus  tomcod  (Walbaum) 196 

Haddock  Melanogrammus  aeglefinus  (Linnaeus) 199 

American  pollock  Pollachius  virens  (Linnaeus) 213 

White  hake  Urophycis  tenuis  (Mitchill) 221 

Squirrel  hake  Urophycis  chuss  (Walbaum) 223 

Spotted  hake  Urophycis  regius  (Walbaum) 230 

Long-finned  hake  Urophycis  chesteri  (Goode  and  Bean)  — 232 

Blue  hake  Antimora  rostrata  Giinther 233 

Hakeling  Physiculus  fulvus  Bean 233 

Four-bearded  rockling  Enchelyopus  cimbrius  (Linnaeus)  — 234 

Cusk  Brosme  brosme  (Miiller) 238 

Grenadiers.     Family  Macrouridae 243 

Common  grenadier  Macrourus  bairdii  Goode  and  Bean 243 

Rough-headed  grenadier  Macrourus  berglax  Lacepede 245 

Long-nosed  grenadier  Coelorhynchus  carminatus  (Goode) 246 

Opahs.     Family  Lampridae 247 

Opah  Lampris  regius  (Bonnaterre) 247 

Flounders   and   Soles.     Families  Hippoglossidae,  Paralichthyidae,  Pleuronectidae, 

Bothidae,  and  Achiridae 248 

Halibut  Hippoglossus  hippoglossus  (Linnaeus) 249 

Greenland  halibut  Reinhardtius  hippoglossoides  (Walbaum) 258 

American  dab  Hippoglossoides  platessoides  (Fabricius) 259 

Summer  flounder  Paralichthys  dentatus  (Linnaeus) 267 

Four-spotted  flounder  Paralichthys  oblongus  (Mitchill) 270 

Yellow-tail  Limanda  ferruginea  (Storer) 271 

Winter  flounder  Pseudopleuronectes  americanus  (Walbaum) 276 

Smooth  flounder  Liopsetta  putnami  (Gill) 283 

Witch  flounder  Glypiocephalus  cynoglossus  (Linnaeus) 285 

Sand  flounder  Lophopsetta  maculala  (Mitchill) 290 

Gulf  Stream  flounder  Citharichthys  arctifrons  Goode 294 

Hogchoker  Achirus  fascialus  Lacepede 296 

John  Dories.     Family  Zeidae 297 

American  John  Dory  Zenopsis  ocellata  (Storer) 297 

Grammicolepid  fishes.     Family  Grammicolepidae 299 

Grammicolepid  Xenolepidichthys  americanus  Nichols  and  Firth 299 

Snipe  fishes.     Family  Macrorhamphosidae 301 

Snipe  fish  Macrorhamphosus  scolopax  (Linnaeus) 301 


VI  CONTENTS 

Bony  fishes.     Class  Osteichthyes — Continued  Page 

Silversides.     Family  Atherinidae 302 

Sil verside  Menidia  menidia  (Linnaeus) 302 

Waxen  silverside  Menidia  beryllina  (Cope) 304 

Mullets.     Family  Mugilidae 305 

Mullet  Mugil  cephalus  Linnaeus 305 

Barracudas.     Family  Sphyraenidae 306 

Northern  barracuda  Sphyraena  borealis  DeKay 306 

Sticklebacks.     Family  Gasterosteidae 307 

Nine-spined  stickleback  Pungitius  -pungitius  (Linnaeus) 307 

Three-spined  stickleback  Gasterosteus  aculeatus  Linnaeus 308 

Two-spined  stickleback  Gasterosteus  wheatlandi  Putnam 310 

Four-spined  stickleback  Apeltes  quadracus  (Mitchill) 311 

Pipefishes.     Family  Syngnathidae 312 

Pipefish  Syngnathus  fuscus  Storer 312 

Pelagic  pipefish  Syngnathus  pelagicus  Linnaeus 314 

Seahorses.     Family  Hippocampidae 315 

Sea  horse  Hippocampus  hudsonius  DeKay .  315 

Trumpetfishes.     Family  Fistulariidae 316 

Trumpetfish  Fistularia  tabacaria  Linnaeus 316 

Mackerels.     Family  Scombridae 317 

Mackerel  Scomber  scombrus  Linnaeus 317 

Chub  mackerel  Pneumatophorus  colias  (Gmelin) 333 

Striped  bonito  Euthynnus  pelamis  (Linnaeus) 335 

False  albacore  Euthynnus  alleteratus  (Rafinesque) 336 

Common  bonito  Sarda  sarda  (Bloch) 337 

Tuna  Thunnus  thynnus  (Linnaeus) 338 

Spanish  mackerel  Scomberomorus  maculatus  (Mitchill) 347 

King  mackerel  Scomberomorus  regalis  (Bloch) 348 

Cavalla  Scomberomorus  cavalla  (Cuvier) 349 

Escolars.     Family  Gempylidae 349 

Escolar  Ruvetius  pretiosus  Cocco 349 

Cutlassfishes.     Family  Trichiuridae 350 

Cutlassfish  Trichiurus  lepturus  Linnaeus 350 

Swordfishes.     Family  Xiphiidae 351 

Swordfish  Xiphias  gladius  Linnaeus 351 

Spearfishes  or  Marlins  and  Sailfishes.     Family  Istiophoridae 357 

Blue  marlin  Makaira  ampla  (Poey) 358 

White  marlin  Makaira  albida  (Poey) 360 

Dolphins.     Family  Coryphaenidae 360 

Dolphin  Coryphaena  hippurus  Linnaeus 360 

Seabreams  or  pomfrets.     Family  Bramidae 361 

Johnson's  Sea  bream  Taractes  princeps  (Johnson) 361 

Butterfishes.     Family  Stromateidae 363 

Butterfish  Poronotus  triacanthus  (Peck) 363 

Harvestfish  Peprilus  alepidotus  (Linnaeus) 368 

Rudderfishes.     Family  Centrolophidae 369 

Barrelfish  Palinurichthys  perciformis   (Mitchill) 369 

Black  ruff  Centrolophus  niger  (Gmelin) 370 

Pompanos  and  Jacks.     Family  Carangidae 371 

Pilotfish  Naucrales  ductor  (Linnaeus) 372 

Rudderfish  Seriola  zonata  (Mitchill) 373 

Mackerel  scad  Decapterus  macarellus  (Cuvier  and  Valenciennes) 374 

Crevalle  Caranx  hippos  (Linnaeus) 375 

Hardtail  Caranx  crysos  (Mitchill) 376 

Saurel  Trachurus  trachurus  (Linnaeus) 377 

Goggle-eyed  scad  Trachurops  crumenopthalmus  (Bloch) 377 

Moonfish  Vomer  setapinnis  (Mitchill) 378 

Lookdown  Selene  vomer  (Linnaeus) 279 

Leatherjacket  Oligoplites  saurus  (Bloch  and  Schneider) 380 

Threadfin  Alectis  crinitus  (MitcbilO 381 


CONTENTS  VII 

Bony  fishes.     Class  Osteichthyes — Continued  P»w 

Bluefishes.     Family  Pomatomidae 382 

Bluefish  Pomatomus  saltalrix  (Linnaeus) 383 

Sea  basses.     Family  Serranidae 389 

Striped  bass  Roccus  saxatilis  (Walbaum) 389 

White  perch  Morone  americana  (Gmelin) 405 

Sea  bass  Centropristes  striatus  (Linnaeus) 407 

Wreckfish  Polyprion  americanus  (Bloch  and  Schneider) 409 

Catalufas  or  Big  Eyes.     Family  Priacanthidae 410 

Short  big-eye  Pseudopriacanlhus  altus  (Gill) 410 

Porgies.     Family  Sparidae 411 

Scup  Stenotomus  versicolor  (Mitchill) 411 

Sheepshead  Archosargus  probatocephalus  (Walbaum) 416 

Croakers,  Drums,  and  Weakfishes.      Family  Sciaenidae 417 

Weakfish  Cynoscion  regalis  (Bloch  and  Schneider) 417 

Spot  Leiostomus  xanthurus  Lacfipede 423 

Kingfish  Menticirrhus  saxatilis  ( Bloch  and  Schneider) 423 

Black  drum  Pogonias  cromis  (Linnaeus) 425 

Tilefishes.     Family  Branchiostegidae 426 

Tilefish  Lopholalilus  chamaeleonticeps  Goode  and  Bean 426 

Rockfishes.     Family  Scorpaenidae 430 

Rosefish  Sebastes  marinus  (Linnaeus) 430 

Black-bellied  rosefish  Helicolenus  dactyloplerus  (De  La  Roche) 437 

Boarfishes.     Family  Caproidae 438 

Boarfish  Antigonia  capros  Lowe 438 

Sculpins  and  Sea  Ravens.     Families  Cottidae  and  Hemitripteridae 439 

Hook-eared  sculpin  Arlediellus  uncinatus  (Reinhardt) 440 

Mailed  sculpin  Triglops  ommatislius  Gilbert 441 

Grubby  Myoxocephalus  aeneus  (Mitchill) 443 

Shorthorn  sculpin  Myoxocephalus  scorpius  (Linnaeus) 445 

Longhorn  sculpin  Myoxocephalus  octodecemspinosus  (Mitchill) 449 

Staghorn  sculpin  Gymnocanthus  tricuspis  (Reinhardt) 452 

Arctic  sculpin  Cottunculus  microps  Collett 453 

Sea  raven  Hemitriplerus  americanus  (Gmelin) 454 

Alligatorfishes.     Family  Agonidae 457 

Alligatorfish  Aspidophoroides  monopterygius  (Bloch) 457 

Lumpfishes.     Family  Cyclopteridae 459 

Lumpfish  Cyclopterus  lumpus  Linnaeus 459 

Spiny  lumpfish  Eumicrotremus  spinosus  (MtiUer) 463 

Sea  snails.     Family  Liparidae 464 

Sea  snail  Neoliparis  atlanticus  Jordan  and  Evermann 464 

Striped  sea  snail  Liparis  liparis  (Linnaeus) 466 

Sea  robins  or  Gurnards  and  Armored  sea  robins.   Families  Triglidae  and  Peristediidae—  467 

Common  sea  robin  Prionotus  carolinus  (Linnaeus) 467 

Striped  sea  robin  Prionotus  evolans  (Linnaeus) 470 

Armored  sea  robin  Peristedion  minialum  Goode 471 

Flying  gurnards.     Family  Dactylopteridae 472 

Flying  gurnard  Dactyloplerus  volitans  (Linnaeus) 472 

Cunner  Tribe  or  Wrasses.     Family  Labridae 473 

Cunner  Tautogolabrus  adspersus  (Walbaum) 473 

Tautog  Tautoga  onitis  (Linnaeus) 478 

Remoras.     Family  Echeneidae 484 

Shark  sucker  Echeneis  naucrates  Linnaeus 485 

Swordfish  sucker  Remora  brachyptera  (Lowe) 486 

Remora  Remora  remora  (Linnaeus) 487 

Sand  launces.     Family  Ammodytidae 487 

Sand  launce  Ammodytes  americanus  DeKay 488 


VIII  CONTENTS 

Bony  fishes.     Class  Osteichthyes— Continued  Fag» 

Blenny-like  fishes.     Families  Lumpenidae,  Pholidae,  and  Stichaeidae 491 

Rock  eel  Pholis  gunnellus  (Linnaeus) 492 

Snake  blenny  Lumpenus  lumpretaeformis  ( Walbaum) 494 

Shanny  Leptoclinus  maculatus  (Fries) 497 

Arctic  shanny  Stichaeus  punctaius  (Fabricius) 497 

Radiated  shanny  Ulvaria  subbifurcala  (Storer) 498 

Wrymouths.     Family  Cryptacanthodidae 500 

Wry  mouth  Cryplacanthod.es  maculatus  Storer 500 

WolfBshes.     Family  Anarhichadidae 502 

WolfBsh  Anarhichas  lupus  Linnaeus 503 

Spotted  WolfHsh  Anarhichas  minor  Olafsen - 507 

Ocean  pouts  and  Wolf  eels.     Family  Zoarcidae 508 

Ocean  pout  Macrozoarces  americanus  (Bloch  and  Schneider) 510 

Wolf  eel  Lycenchelys  verrillii  (Goode  and  Bean) 515 

Arctic  eelpout  Lycodes  reticulalus  Reinhardt 516 

Cusk  eels.     Family  Ophidiidae 517 

Cusk  eel  Lepophidium  cervinum  (Goode  and  Bean) 517 

Toadfishes.     Family  Batrachoididae 518 

Toadfish  Opsanus  tau  (Linnaeus) 518 

Triggerfishes.     Family  Balistidae 520 

Triggerfish  Batistes  carolinensis  Gmelin 520 

Filefishes.     Family  Monacanthidae 521 

Filefish  Monacanthus  hispidus  (Linnaeus) 522 

Filefish  Monacanthus  ciliatus  (Mitchill) 523 

Orange  filefish  Alutera  schoepfii  (Walbaum) 524 

Unicornfish  Alutera  scripta  (Gmelin) 525 

Puffers  and  Porcupine-fishes.     Families  Tetraodontidae  and  Diodontidae 525 

Puffer  Sphaeroides  maculatus  (Bloch  and  Schneider) 526 

Burrfish  Chilomycterus  schoepfii  (Walbaum) 527 

Ocean  Sunfishes  or  Headfishes.     Family  Molidae 528 

Sunfish  Mola  mola  (Linnaeus) 529 

Sharp-tailed  sunfish  Masturus  lanceolatus  (Lifinard) 53 1 

Anglers.     Family  Lophiidae 532 

American  Goosefish  Lophius  americanus  Cuvier  and  Valenciennes 532 

Sargassum  fishes.     Family  Antennariidae 541 

Sargassum  fish  Histrio  piclus  (Cuvier  and  Valenciennes) 541 

Deep  sea  anglers.     Family  Ceratiidae 542 

Deep  sea  angler  Ceratias  holbolli  Kr0yer 543 

Bibliography 545 

Index 561 


FISHES  OF  THE  GULF  OF  MAINE 

By  HENRY  B.  BIGELOW  and  WILLIAM  C.  SCHROEDER 
Museum  of  Comparative  Zoology,  Harvard  University,  and  Woods  Hole  Oceanographic  Institution 


During  the  summer  of  1912  the  Bureau  of 
Fisheries,  with  the  cooperation  of  the  Museum 
of  Comparative  Zoology  of  Harvard  University, 
commenced  an  oceanographic  and  biological 
survey  of  the  Gulf  of  Maine,  with  special  refer- 
ence to  its  fishes,  to  its  floating  plants  and  animals 
(plankton),  to  the  physical  and  chemical  state  of 
its  waters,  and  to  the  circulation  of  the  latter. 
Cruises  were  made  on  the  Fisheries  schooner 
Grampus  during  the  summers  and  autumns  of 
1912,  1913,  1914,  1915  and  1916,  and  during  the 
winters  and  springs  of  1913  and  1915.  The  work 
was  interrupted  by  the  war,  but  was  resumed  with 
a  cruise  of  the  Fisheries  steamer  Albatross  in  the 
late  winter  and  spring  of  1920,  and  was  continued 
by  the  Fisheries  steamer  Halcyon  during  the 
winter  and  spring  of  1920-21,  and  the  summers 
of  1921  and  1922. 

The  first  part  of  the  general  report,  dealing 
with  the  fishes,  was  published  in  1925,  as  Bulletin 
40  (Pt.  1)  of  the  United  States  Bureau  of  Fisher- 
ies ; '  subsequent  parts  describing  the  plankton  of 
the  offshore  waters  of  the  Gulf  and  the  physical 
characteristics  of  its  waters  were  published  in 
1926-27,  as  Part  2. 

The  preparation  of  the  section  on  the  fishes  was 
assigned  originally  to  W.  W.  Welsh,  who  had 
gathered  a  large  body  of  original  observations  on 
the  growth,  reproduction,  diet,  and  other  phases 
of  the  lives  of  many  of  the  more  important  species. 
The  report  was  far  advanced  when  it  was  inter- 
rupted by  his  untimely  death,  and  H.  B.  Bigelow 
undertook  to  carry  it  to  publication  along  the 
lines  originally  laid  down.  The  new  edition, 
entailing  a  general  revision  and  the  addition  of 
much  new  material,  has  been  prepared  jointly  by 
H.  B.  Bigelow  and  by  W.  C.  Schroeder. 

i  The  Bureau  of  Fisheries  was  transferred  on  July  1, 1939,  from  the  Depart- 
ment of  Commerce  to  the  Department  of  the  Interior,  and  on  July  30,  1940, 
it  was  consolidated  with  the  Bureau  of  Biological  Survey  to  form  the  Fish 
and  Wildlife  Service. 

210941—53 2 


AREA  COVERED 

The  term  "Gulf  of  Maine"  covers  the  oceanic 
bight  from  Nantucket  Shoals  and  Cape  Cod  on 
the  west,  to  Cape  Sable  on  the  east.  Thus  it 
includes  the  shore  lines  of  northern  Massachu- 
setts, New  Hampshire,  Maine,  and  parts  of  New 
Brunswick  and  of  Nova  Scotia.  The  eastern 
and  western  boundaries  adopted  in  this  paper  are 
65°  and  70°  West  longitude,  respectively.  South- 
ern strays,  or  northern,  which  have  no  real  status 
in  the  Gulf  of  Maine  except  by  accident,  are 
mentioned  only  briefly,  or  are  relegated  to  foot- 
notes. The  Gulf  of  Maine  has  a  natural  seaward 
rim  formed  by  Nantucket  Shoals,  by  Georges 
Bank,  and  by  Browns  Bank.  We  have  chosen 
the  150-fathom  contour  as  the  arbitrary  offshore 
boundary,  because  this  will  include  all  of  the 
species  that  are  likely  to  be  caught  by  commer- 
cial fishermen  but  will  exclude  almost  the  entire 
category  of  the  so-called  "deep-sea"  fishes,  which 
are  numerous  in  the  basin  of  the  open  Atlantic 
but  are  not  constituents  of  the  fauna  of  the  Gulf 
of  Maine,  properly  speaking. 

The  general  oceanography  of  this  area  has  been 
the  subject  of  another  report,  but  it  may  not  be 
amiss  to  point  out  that  the  temperature  of  the 
Gulf  and  its  fauna  are  boreal,  and  that  its  south- 
ern and  western  boundaries  are  the  northern 
limit  to  common  occurrence  of  many  southern 
species  of  fishes  and  of  invertebrates. 

SCOPE  OF  THE  WORK 

Our  aim  has  been  a  handbook  for  the  easy 
identification  of  the  fishes  that  occur  in  the  Gulf 
of  Maine,  with  summaries  of  what  is  known  of 
the  distribution,  relative  abundance,  and  more 
significant  facts  in  the  life  history  of  each.  The 
descriptions  are  as  little  technical  as  is  com- 
patible with  scientific  accuracy,  and  are  limited 

1 


FISHERY   BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


chiefly  to  such  of  the  external  features  of  each 
kind  of  fish  as  may  serve  for  identification  in  the 
field. 

References  to  more  detailed  descriptions  and 
synonymies  are  given  to  Bigelow  and  Schroeder 
(Fishes  of  the  Western  North  Atlantic,  Parts  1 
and  2,  1948,  1953)  for  the  cartilaginous  fishes;  also 
to  Garman's  beautiful  plates  for  such  of  these  as 
he  pictured  in  his  classic  monograph,  published  in 
1913,  in  vol.  36,  of  the  Memoirs  of  the  Museum  of 
Comparative  Zoology.  References  for  the  various 
species  of  bony  fishes  are  to  Jordan  and  Ever- 
mann's  Fishes  of  North  and  Middle  America 
(Bulletin  47,  U.  S.  National  Museum,  1896-1900, 
Parts  1—4),  which  still  remains  the  only  compre- 
hensive work  on  the  bony  fishes  of  North  America. 
Many  of  the  illustrations  have  been  borrowed 
from  earlier  publications,  but  some  of  them  are 
original. 

Keys  are  provided  for  all  species  as  a  further 
aid  to  identification. 

In  most  cases  the  sizes  of  larval  fish  and  eggs  are 
given  in  millimeters  (1  inch  equals  25.4  mm.);  the 
sizes  of  the  larger  fishes  are  in  inches  and  feet; 
weights  are  in  pounds. 

The  scientific  nomenclature  of  the  cyclostomes, 
of  the  elasmobranchs,  and  of  the  chimaeroids, 
follows  Bigelow  and  Schroeder  (Fishes  of  the 
Western  North  Atlantic,  No.  1,  Parts  1  and  2, 
1948;  1953)  that  of  the  bony  fishes  follows  Jordan, 
Evermann,  and  Clark's  Check  List  of  the  Fishes 
and  Fishlike  Vertebrates  of  North  and  Middle 
America  (Report,  U.  S.  Commissioner  of  Fisheries 
for  1928  (1930),  Part  2),  unless  otherwise  noted. 
The  families  of  bony  fishes  are  arranged  for  the 
most  part  in  the  sequence  employed  by  Jordan, 
Evermann,  and  Clark,  except  that  the  several 
families  of  luminescent  fishes  are  grouped  together, 
in  the  hope  of  making  it  easier  for  the  nontechnical 
observer  to  identify  such  of  them  as  may  come  to 
hand. 

SOURCES  OF  INFORMATION 

The  literature  dealing  with  the  fishes  of  the 
Gulf  of  Maine  begins  with  the  earliest  descriptions 
of  New  England.  Captain  John  Smith,  for  in- 
stance, commented  on  the  abundance  of  sturgeon, 
cod,  hake,  haddock,  cole  (the  American  pollock), 
cusks,  sharks,  mackerel,  herring,  cunners,  eels, 
salmon,  and  striped  bass,  in  his  Generall  Historie 
of  Virginia,  New  England  and  the  Summer  Isles, 


published  in  1616,  while  Wood  in  his  New  Eng- 
land's Prospect,  1634,  gave  much  interesting  infor- 
mation, some  of  which  we  quote  hereafter. 

The  sea  fishes  of  northern  New  England  and  of 
the  Maritime  Provinces  had  begun  to  attract 
scientific  attention  by  the  early  part  of  the  nine- 
teenth century,  and  many  local  faunal  lists  have 
been  published  since  then.  The  following  are  the 
most  important  of  these,  in  chronological  arrange- 
ment: 

1850.  Report  on  the  sea  and  river  fisheries  of  New 
Brunswick,  within  the  Gulf  of  St.  Lawrence  and  Bay  of 
Chaleur,  M.  H.  Perley,  137  pp.,  1850.  Fredericton,  New 
Brunswick. 

1853-1867.  A  history  of  the  fishes  of  Massachusetts, 
David  Humphreys  Storer.  Memoirs,  American  Academy 
of  Arts  and  Sciences,  New  Series,  vol.  5,  pp.  49-92,  122-168, 
and  257-296;  vol.  6,  pp.  309-372;  vol.  8,  pp.  389-439;  vol. 
9,  pp.  217-256,  39  pis.  (Also  in  book  form  with  supple- 
ment, 1867),  Cambridge  and  Boston. 

1879.  A  list  of  the  fishes  of  Essex  County,  including 
those  of  Massachusetts  Bay,  George  Brown  Goode,  and 
Tarleton  H.  Bean.  Bulletin,  Essex  Institute,  vol.  11,  No. 
1,  pp.  1-38.     Salem. 

1884.  Natural  history  of  useful  aquatic  animals,  George 
Brown  Goode  and  associates,  Section  I,  The  Fisheries  and 
Fishery  Industries  of  the  United  States.  Published  jointly 
by  the  U.  S.  Fish  Commission  and  the  U.  S.  Bureau  of  the 
Census,  895  pp.     Washington. 

1908.  Fauna  of  New  England.  8.  List  of  the  Pisces, 
William  C.  Kendall.  Occasional  Papers,  Boston  Society 
of  Natural  History,  vol.  7,  No.  8,  April  1908,  pp.  1-52. 
Boston. 

1914.  An  annotated  catalogue  of  the  fishes  of  Maine, 
William  C.  Kendall.  Proceedings,  Portland  Society  of 
Natural  History,  vol.  3,  1914,  Part  1,  pp.  1-198.     Portland. 

1922.  The  fishes  of  the  Bay  of  Fundy,  A.  G.  Huntsman. 
Contributions  to  Canadian  Biology  (1921),  1922,  No.  3, 
pp.   1-24  (51-72).     Ottawa. 

These  lists  contain  all  the  early  published  local- 
ity records  of  the  rarer  species,  either  first  hand, 
or  by  reference  to  original  sources,  while  the  last 
two,  with  a  paper  by  Gill,2  and  the  first  edition  of 
the  present  book  give  complete  bibliographies 
for  the  Canadian  coasts  of  the  Gulf  and  for  the 
coasts  of  Maine  and  of  Massachusetts.  A  similar 
list  of  the  captures  of  deep  water  fishes  along  the 
outer  part  of  the  continental  shelf  is  to  be  found  in 
Goode  and  Bean's   "Oceanic  Ichthyology." 3 

The  most  pertinent  extralimital  lists  are  Smith's4 
and  Sumner,  Osburn  and  Cole's  6  lists  of  Woods 


>  Kept.  U.  S.  Comm.  Fish.,  (1904)  1905,  pp.  163-188. 
i  Smithsonian  Contribution  to  Knowledge,  vol.  30, 1895. 
'  Bull.  U.  S.  Fish  Comm.,  Vol.  17, 1898,  pp.  85-111. 
'  Bull.  U.  S.  Bur.  Fish.,  vol.  31,  Pt.  2, 1913,  pp.  549-794. 


FISHES   OF  THE  GULF  OF   MAINE 


Hole  fishes;  Halket's  8  Checklist  of  the  fishes  of 
Canada  and  of  Newfoundland,  and  Vladykov  and 
McKenzie's  The  Marine  Fishes  of  Nova  Scotia.7 

The  literature  dealing  with  the  habits  of  the 
fishes  of  the  Gulf  of  Maine  is  very  extensive,  for 
most  of  the  important  commercial  species,  and 
many  of  the  others  also,  are  common  to  both  sides 
of  the  North  Atlantic.  Among  general  European 
manuals,  Day's  Fishes  of  Great  Britain  and  Ire- 
land,8 Smitt's  "Scandinavian  Fishes,"  9  and  Ehren- 
baum's  summary  of  the  many  scattered  accounts 
of  the  eggs  and  larvae  of  northern  fishes10  have 
been  especially  helpful. 

A  large  amount  of  information  as  to  local  dis- 
tribution and  abundance  of  various  fishes  has  been 
gleaned  from  unpublished  material  in  the  files  of 
the  U.  S.  Fish  and  Wildlife  Service,  as  well  as  from 
the  fishery  statistics  published  by  the  Fisheries 
Branch,  U.  S.  Fish  and  Wildlife  Service  (formerly 
the  U.  S.  Bureau  of  Fisheries),  by  the  Dominion  of 
Canada,  and  by  the  Commonwealth  of  Massa- 
chusetts. The  superintendents  of  the  Woods 
Hole,  Gloucester,  and  Boothbay  hatcheries  have 
supplied  much  valuable  information,  as  have  other 
members  of  the  U.  S.  Fish  and  Wildlife  Service. 
Among  these,  Leslie  Scattergood  has  given  many 
interesting  pieces  of  information  for  Maine  waters, 
while  Howard  Schuck  has  contributed  authentic- 
ity to  the  account  of  the  haddock.  Dr.  A.  G. 
Huntsman  has  contributed  his  unpublished  notes 
on  the  fishes  of  the  Bay  of  Fundy  and  Gulf  of 
St.  Lawrence.  Dr.  A.  H.  Leim,  Mr.  R.  A.  McKen- 
zie,  and  Dr.  Vadim  D.  Vladykov  have  supplied  us 
with  pertinent  information  on  certain  species  from 
the  Nova  Scotian-St.  Lawrence  River  regions. 
The  late  Prof.  J.  P.  McMurrich  permitted  the 
use  of  his  unpublished  plankton  records,  and  a 
number  of  Newfoundland  records  were  furnished 
by  Drs.  George  W.  Jeffers  and  E.  Templeman. 

The  late  W.  F.  Clapp  has  contributed  many 
interesting  notes  gleaned  during  his  experience  as 
a  fisherman  before  entering  the  scientific  field. 
Harry  Piers  of  the  Provincial  Museum  of  Halifax, 

•  Checklist  of  the  Fishes  ot  the  Dominion  of  Canada  and  Newfoundland 
1913,  138  pp. 

»  Proc.  Nova  Scotia  Inst,  of  Science,  vol.  19,  Pt.  1, 1935,  pp.  17-113. 

»  The  fishes  of  Great  Britain  and  Ireland,  by  F.  Day,  Text  vol.  1,  CXII+ 
336  pp.,  vol.  2,  388  pp.,  and  atlas,  179  plates,  1880-1884.  London  and  Edin- 
burgh. 

•  A  history  of  Scandinavian  fishes.  Second  edition,  vol.  1,  1892;  vol.  2, 
1895;  1,240  pp.,  53  pis.    Stockholm. 

10  Eier  und  Larven  von  Fischen.  Nordisches  Plankton,  vol.  I,  413  pp.,  148 
figs.;  appeared  in  two  parts  as  Lief.  4, 1905,  and  Lief.  10, 1919. 


has  supplied  interesting  information  on  the  occur- 
rence of  the  blue  shark.  John  Worthington  has 
furnished  us  with  pound-records  for  the  Truro- 
Provincetown  region  covering  a  recent  span  of 
about  fifteen  years  and  has  given  us  specimens  of 
three  species  heretofore  unreported  in  the  Gulf  of 
Maine.  Benjamin  H.  Morrow  has  supplied  inter- 
esting data  from  the  vicinity  of  Sandwich,  Mass. 
We  have  received  much  information  about  the 
striped  bass  in  Nova  Scotia  from  Major  Howard 
Scott,  through  the  kind  offices  of  Henry  Lyman. 
And  we  owe  it  to  consultation  with  Dr.  A.  Vedel 
Tuning;  of  the  Marine  Biological  Laboratory, 
Charlottenlund,  Denmark,  and  the  specimens 
contributed  by  Dr.  C.  E.  Lucas  of  the  Scottish 
Fisheries  Laboratory,  Aberdeen,  that  we  have 
dared  to  reach  a  conclusion  as  to  the  relationship 
between  the  rosefish  of  our  gulf  and  of  north 
European  waters.  Francis  Sargent,  also  of  the 
Division  of  Marine  Fisheries  of  Massachusetts, 
and  Henry  Lyman,  editor  of  the  Salt  Water 
Sportsman,  have  been  unfailing  in  their  response 
to  our  many  inquiries.  Myvanwy  Dick  of  the 
Harvard  Museum  of  Comparative  Zoology  has 
been  of  assistance  in  the  handling  of  certain  of  our 
study  material  and  in  the  preparation  of  a  number 
of  illustrations.  The  illustrations  of  the  hagfish 
and  lamprey  and  most  of  those  of  the  sharks, 
skates,  rays,  and  chimaera  are  reprinted  here 
through  the  courtesy  of  the  Sears  Foundation  for 
Marine  Research,  publisher  of  the  Fishes  of  the 
Western  Atlantic,  Memoir  1,  Parts  1  and  2,  in 
which  the  illustrations  originally  appeared.  Claude 
Ronne  of  the  Woods  Hole  Oceanographic  Institu- 
tion prepared  many  photographs  from  both 
original  and  published  drawings,  which  were  used 
to  illustrate  this  book. 

We  owe  a  debt  of  gratitude,  also,  to  the  late 
Dr.  Samuel  Garman,  who  was  ever  ready  with 
assistance  until  the  time  of  his  death,  and  to 
W.  C.  Adams,  former  director  of  the  division  of 
fisheries  and  game  of  the  State  of  Massachusetts. 
We  wish  to  express  our  hearty  thanks  to  the  many 
commercial  fishermen  and  to  the  many  salt  water 
anglers  of  our  acquaintance  who  have  met  our 
inquiries  in  the  most  cordial  way  and  who  have 
supplied  us  with  a  vast  amount  of  first-hand 
information  on  the  habits,  distribution,  and 
abundance  of  the  commercial  and  game  fishes, 
which  could  be  had  from  no  other  source.     The 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


preparation  of  this  book  would  have  been  out  of 
the  question  without  their  help. 

Finally,  we  have  ourselves  gathered  a  large 
body  of  data  as  to  distribution,  habits,  spawning 
seasons,  and  like  matters,  through  many  years, 
at  many  localities,  both  inshore  and  on  the  offshore 
banks. 

USE  OF  THE  KEYS 

The  various  fins  and  other  structures  mentioned 
in  the  keys  are  named  in  the  accompanying  out- 
lines of  a  haddock  and  of  a  typical  shark  (fig.  1). 
A  simple  way  to  explain  the  use  of  the  keys  is  to 
use  the  haddock  as  an  example,  running  it  down 
with  the  illustration  at  hand  for  reference. 

Turning  to  Key  A.  (p.  5),  we  find  that  our  fish 
fits  the  second  alternative  under  section  1,  since 
it  has  bony  jaws  and  pectoral  fins,  and  is  not 
shaped  like  an  eel.     This  refers  us  to  section  3. 

There  being  only  one  gill  opening  on  each  side, 
we  go  from  section  3  to  section  5.  As  our  fish 
does  not  have  a  tubular  snout  section  5  refers  us 
to  section  6,  and  this  in  turn  to  section  7,  since 
neither  the  upper  jaw  nor  the  lower  is  greatly 

1st  Dorsal  Fin 


prolonged.  Since  the  body  is  not  square-cut  close 
behind  the  dorsal  and  anal  fins,  but  has  a  definite 
tail  part,  we  proceed  from  section  7  to  section  8, 
and  from  section  8  to  section  11,  for  our  fiah  has 
no  sucking  plate  or  disc,  either  on  top  of  the  head, 
or  on  the  chest.  Section  11  refers  us  in  turn  to 
section  12  because  the  tail  fin  is  nearly  symmetri- 
cal in  outline.  The  anal  fin  being  clearly  and 
definitely  separated  from  the  caudal  fin,  we  go 
from  section  12  to  section  13;  and  from  section  13 
to  section  14,  for  our  fish  does  not  have  any  evi- 
dent light-producing  ("luminescent")  spots  either 
on  its  sides  or  on  its  head.  Our  fish  does  not 
have  a  fleshy  fin  or  flap  either  in  front  of  the  ordi- 
nary dorsal  fins  or  behind  them,  but  all  of  its 
dorsal  fins  are  supported  by  rays  that  are  visible  if 
held  against  the  light.  Consequently,  we  proceed 
from  section  14  to  section  18,  and  this  refers  us 
to  section  22,  there  being  no  flaps  or  tags  of  skin 
on  the  sides  of  the  head.11  Our  fish  obviously 
does  not  lie  flat  on  one  side,  i.  e.,  it  is  not  one  of 
the  flat  fishes,  which  brings  us  to  section  23,  and 

i'  There  Is  a  barbel  on  its  chin,  but  this  Is  very  different  In  appearance  from 
the  skin  flaps  around  the  jaws  that  are  characteristic  of  the  few  species  that 
fall  under  the  first  alternative  of  section  IS. 


2nd  Donal 
Fin  i 


Procoudal 
Pit 


Caudal 


tst.Ana.1  fin 
Ventral  fin 

Figure  1. — Diagrams  of  a  haddock  (below)  and  of  a  typical  shark  (above)  with  terms  used  in  the  keys  and  descriptions. 


FISHES   OF  THE  GULF   OF   MAINE  5 

this  in   turn  carries  us  to  Key  E  (p.  7)  because  which  refers  it  to  section  2.     And  here  the  black 

it  has  three  separate,  well  developed  dorsal  fins.  lateral  line  and  the  dark  blotch  on  each  shoulder 

Since  there  are  3  dorsal  fins  and  2  anal  fins,  name  it  a  haddock, 
section  1  of  Key  E  sends  us  to  the  key  to  the  cod  Any  other  Gulf  of  Maine  species  is  to  be  named 

and  silver  hake  families  (p.  173).  Turning  to  the  in  the  same  way,  starting  with  Key  A,  section  1, 

first  section  of  the  latter  we  find  that  our  fish  fits  and  following  through  the  appropriate  alternatives 

the  first  alternative  (3  dorsal  fins  and  2  anals) ,  as  thev  refer  it  from  section  to  section. 

KEY  TO  GULF  OF  MAINE  FISHES 
Key  A 

1.  Mouth  soft,  with  no  firm  jaws;  no  pectoral  fins;  form  eel-like 2 

Mouth  has  firm  jaws;  pectoral  fins  are  present  even  if  the  form  is  eel-like 3 

2.  Two  separate  fins  on  the  back;  no  barbels  on  the  snout Lamprey,  p.  12 

Only  one  fin  on  the  back;  with  barbels  on  the  snout Hag,  p.  10 

3.  Five  gill  openings  on  each  side 4 

Only  one  gill  opening  on  each  side 5 

4.  General  form  cylindrical  in  all  Gulf  of  Maine  species;  the  forward  edges  of  the  pectoral  fins  are  not  attached  to  the 

sides  of  the  head  forward,  past  the  gill  openings;  the  gill  openings  are  not  confined  to  the  lower  surface;  the  upper 

margin  of  each  orbit  is  free  from  the  eyeball,  as  a  free  eyelid Sharks,  key,  p.  16 

General  form  very  fiat,  disclike;  the  forward  edges  of  the  pectoral  fins  are  attached  to  the  sides  of  the  head  forward 
past  all  of  the  gill  openings;  the  gill  openings  are  confined  to  the  lower  surface;  the  upper  margin  of  each  orbit  is 
not  free  from  the  eyeball  (no  free  eyelid) Skates  and  Rays,  key,  p.  57 

5.  The  bones  of  the  head  are  fused  in  a  tubular  snout,  with  the  mouth  at  its  lip refer  to  Key  B,  p.  6 

No  tubular  snout 6 

6.  One  or  both  jaws  are  prolonged  as  a  bony  sword  or  bill _ refer  to  Key  C,  p.  6 

Neither  jaw  is  greatly  prolonged 7 

7.  Body  abruptly  square-cut,  close  behind  the  very  high  dorsal  and  anal  fins refer  to  Sunfishes,  key,  p.  529 

Body  with  distinct  tail  part 8 

8.  There  is  a  sucking  plate  or  disc,  either  on  the  top  of  the  head  or  on  the  chest 9 

There  is  no  sucking  disc  or  plate 11 

9.  The  sucking  plate  is  on  the  top  of  the  head refer  to  Remora  family,  key,  p.  485 

The  sucking  disc  is  on  the  chest 10 

10.  General  form  is  like  a  tadpole;  the  anal  fin  originates  about  as  far  back  as  the  tips  of  the  pectorals. 

refer  to  Sea  snail  family,  key,  p.  464 

General  form  is  not  like  a  tadpole,  but  is  high  arched,  with  longitudinal  ridges;  the  anal  fin  originates  far  behind  the 

tips  of  the  pectorals refer  to  Lumpfish  family,  key,  p.  459 

11.  Tail  like  a  shark,  i.  e.,  with  the  upper  lobe  much  longer  than  the  lower Sturgeons,  key,  p.  81 

Tail  with  the  upper  and  lower  lobes  of  equal  lengths,  or  nearly  so 12 

12.  No  clear  separation  between  the  anal  and  the  caudal  fins,  which  together  form  one  continuous  fin  (the  anal  portion 

may  be  either  long  or  short) refer  to  Key  D,  p.  6 

Anal  and  caudal  fins  are  separated  by  a  deep  notch,  or  by  a  space 13 

13.  Sides  of  body  and  head,  or  both,  with  luminescent  spots  or  patches,  easily  seen  if  not  damaged. 

refer  to  Luminescent  fishes,  key,  p.  141 
No  luminescent  organs 14 

14.  There  is  a  fleshy  ("adipose")  fin,  with  neither  rays  nor  spines,  either  in  front  of  the  rayed  dorsal  fin,  or  behind  it..  15 
There  is  no  fleshy  ("adipose")  fin,  but  both  the  dorsals  (if  there  are  two)  are  supported  by  rays  or  by  spines  that  can 

be  felt,  if  not  seen 18 

15.  The  adipose  fin  is  on  the  nape  of  the  neck,  in  front  of  the  dorsal  fin Tilefish,  p.  426 

The  adipose  fin  is  behind  the  dorsal  fin 16 

16.  The  dorsal  fin  extends  nearly  the  entire  length  of  the  body Lancetfish,  p.  161 

The  dorsal  fin  is  short,  standing  about  midway  of  the  body 17 

17.  Tail  deeply  forked refer  to  Smelts  and  Argentine  Key,  p.  133 

Tail  nearly  square  or  only  slightly  forked refer  to  Salmon  key,  p.  120 

18.  The  head  is  fringed  with  fleshy  tags  or  flaps 19 

The  head  is  not  fringed  with  fleshy  tags  or  flaps 22 

19.  The  pectorals  are  armlike 20 

The  pectorals  are  not  armlike 21 


6  FISHERY  BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 

Key  A — Continued 

20.  Body  very  broad  and  flat;  mouth  enormous Goosefish,  p.  532 

Body  deep  and  flattened  sidewise;  mouth  small Sargassum  fish,  p.  541 

21.  The  first  (spiny)  dorsal  fin  is  longer  than  the  second  (soft-rayed  dorsal) ;  neither  dorsal  fin  is  fleshy Sea  raven,  p.  454 

The  first  (spiny)  dorsal  fin  is  much  shorter  than  the  second  (soft-rayed  dorsal) ;  both  of  the  dorsals  are  thick  and 

fleshy Toadfish,  p.  518 

22.  Fishes  which  lie  flat  on  the  one  side,  with  both  of  their  eyes  on  the  other  side;  the  upper  side  is  dark,  the  lower  side 

normally  is  pale .. refer  to  Flatfish  tribe  key,  p.  248 

Fishes  which  do  not  lie  flat  on  one  side 23 

23.  Two  or  more  separate  and  well-developed  dorsal  fins,  each  with  continuous  membrane refer  to  Key  E,  p.  7 

Only  one  well-developed  dorsal  fin  with  continuous  membrane  (this,  however,  may  be  preceded  by  isolated  spines  or 

rays) 24 

24.  Top  of  snout  with  several  barbels  or  beards Rockling  (cod  family  in  part) ,  p.  234 

No  barbels  or  beards  on  the  top  of  the  snout 25 

25.  Jaws  with  very  large  canine  tusks refer  to  Wolffishes  key,  p.  503 

No  large  canine  tusks  in  either  jaw 26 

26.  Dorsal  fin  soft-rayed  throughout  its  length,  except  that  it  may  be  preceded  by  a  few  separate  spines. 

refer  to  Key  F,  p.  8 
At  least  the  forward  one-third  of  the  dorsal  fin  is  with  stiff  sharp  rays  or  spines refer  to  Key  G,  p.  9 

Key  B 

Fishes  with  tubular  snouts  (from  No.  5,  p.  5). 

1.  Head  is  horselike;  rear  portion  of  trunk  is  slender,  prehensile;  no  caudal  fin Sea  horse,  p.  315 

Head  is  not  horselike;  rear  part  of  trunk  is  not  prehensile;  there  is  a  caudal  fin 2 

2.  Body  and  head  (measured  from  tip  of  snout)  are  only  about  4  times  as  long  as  deep;  the  dorsal  fin  has  a  long,  strong, 

saw-edged  spine Snipefish,  p.  301 

Body  and  head  (measured  from  tip  of  snout)  are  at  least  25  times  as  long  as  deep;  the  dorsal  fin  does  not  have  a 
large  spine 3 

3.  The  snout  is  not  longer  than  the  dorsal  fin;  the  anal  fin  is  very  small;  no  ventral  fins;  the  caudal  fin  is  rounded 

Pipefishes,  key,  p.  312 

The  snout  is  more  than  6  times  as  long  as  the  dorsal  fin;  the  anal  fin  is  about  as  large  as  the  dorsal;  ventral  fins  are 

present  though  small;  the  caudal  fin  is  forked Trumpetfish,  p.  316 

KeyC 

Fishes  with  bills  or  swords  (from  No.  6,  p.  5) 

1.  Both  of  the  jaws  are  elongated 4 

Only  one  of  the  jaws  is  elongated 2 

2.  Upper  jaw  elongated,  as  a  sword 3 

Lower  jaw  elongated.. Halfbeak,  p.  169 

3.  The  sword  is  flattened  dorso-ventrally,  and  is  sharp-edged;  the  first  dorsal  fin  is  shorter  than  the  sword  forward  of 

the  eyes;  no  ventral  fins Swordfish,  p.  351 

The  sword  is  round-edged;  the  first  dorsal  fin  is  nearly  twice  as  long  as  the  sword refer  to  Spearfishes  or  Marlins 

and  Sailfish,  key,"  p.  358 

4.  The  caudal  fin  is  well  developed 5 

No  caudal  fin;  the  tip  of  the  tail  is  whip-like Snipe  eel,  p.  159 

5.  There  are  several  finlets  behind  the  dorsal  and  anal  fins Needlefish,  p.  170 

No  finlets  behind  the  dorsal  and  anal  fins refer  to  Billfishes  or  Silver  gars,  key,  p.  167 

KeyD 

Bony  fishes  with  snouts  of  ordinary  form;  with  only  one  gill  opening  on  each  side,  and  with  the  anal  fin  continuous 
with  the  caudal  fin  around  the  tip  of  the  tail  (from  No.  12,  p.  5). 

1.  Only  one  dorsal  fin 2 

Two  separate  dorsal  fins,  the  first  much  bigger  than  the  second,  but  shorter 7 

2.  Body  band-shaped,  the  tail  tapering  to  a  whip-like  tip Cutlassfish,  p.  350 

Body  thick,  eel-like;  the  vertical  fins  continuous  around  the  tip  of  the  tail  in  a  broad  band 3 

3.  The  dorsal  fin  is  spiny  from  end  to  end 4 

The  dorsal  fin  is  soft-rayed,  at  least  for  almost  all  its  length 5 

■'  The  sallflsh  would  also  come  under  this  heading  should  one  ever  be  taken  In  the  Gulf  of  Maine.     The  distinctions  between  It  and  the  spearfishts  are 
Blven  on  page  358. 


FISHES   OF  THE  GULF  OF   MAINE  7 

Key  D — Continued 

4.  Mouth  large  and  strongly  oblique ;  there  are  no  ventral  fins Wrymouth,  p.  500 

Mouth  small  and  horizontal;  with  small  ventral  fins Rock  eel,  p.  492 

5.  There  are  no  ventral  fins refer  to  Eel  family  key,  p.  150 

With  small  but  distinct  ventral  fins,  situated  forward  of  the  pectorals 6 

6.  The  ventrals  are  situated  behind  the  gill  openings  and  are  of  ordinary  form refer  to  Eelpout  family,  key,  p.  509 

The  ventrals  are  situated  on  the  chin,  well  in  front  of  the  gill  openings  and  are  reduced  to  forked,  barbel-like  struc- 
tures   Cusk  eel,  p.  517 

7.  The  ventral  fins  are  situated  below  the  points  of  origin  of  the  pectorals;  the  skin  is  conspicuously  scaly. 

refer  to  Grenadier  family,  key,  p.  243 
The  ventral  fins  are  situated  far  back,  behind  the  tips  of  the  pectorals;  the  skin  is  soft,  without  scales. 

Chimaera,  p.  79 

Key  E 

Bony  fishes  of  ordinary  form,  with  2  or  3  well-developed  dorsal  fins  and  with  the  anal  fin  and  the  rearmost  dorsal 
separated  from  the  caudal  fin.  (from  No.  23,  p.  6). 

1.  Three  dorsal  fins  and  2  anal  fins refer  to  Cod  family,  key   (in  part),  p.  173 

Only  2  dorsal  fins  and  1  anal  fin 2 

2.  With  one  or  more  small  finlets  between  the  second  dorsal  and  anal  fins  and  the  caudal  fin 3 

No  finlets  between  the  second  dorsal  and  anal  fins  and  the  caudal  fin 4 

3.  With  more  than  3  dorsal  finlets  and  3  anal  finlets refer  to  Mackerel  family,  key,  p.  317 

With  only  2  dorsal  finlets  and  2  anal  finlets Escolar,  p.  349 

With  only  1  dorsal  finlet  and  1  anal  unlet Mackerel  scad  (Pompano  family,  in  part),  p.  374 

4.  Head  very  broad;  top  and  sides  of  head  bony,  with  sharp  spines  or  horns 5 

Head  not  noticeably  broad;  sides  of  head  have  no  spines  or  horns 7 

5.  First  (lower)  few  rays  of  the  pectoral  fins  are  not  separate  from  the  remainder  of  the  fin;  the  mouth  is  very  large 

refer  to  Sculpin  family,  key,  p.  440 

First  (lower)  few  rays  of  the  pectoral  fins  are  separate  from  the  remainder  of  the  fin;  the  mouth  is  not  very 

large 6 

6.  Each  of  the  first  (lower)  2  or  3  rays  of  the  pectoral  fins  have  the  form  of  a  separate  feeler;  outline  of  tip  of  snout  is 

concave;  the  first  few  spines  of  the  first  dorsal  fin  are  not  separate  from  the  remainder  of  the  fin. 

refer  to  Sea  robin  and  Armored  sea  robin,  key,  p.  467 
First  Power)  few  rays  of  the  pectorals  do  not  have  the  form  of  feelers,  but  are  connected,  one  with  the  next,  by  mem- 
brane, as  a  separate  fin;  outline  of  tip  of  snout  convex;  the  first  few  spines  of  the  first  dorsal  are  separate. 

Flying  gurnard,  p.  472 

7.  First  spine  of  first  dorsal  fin  is  very  much  stouter  than  the  other  spines,  and  can  be  locked  erect  by  the  second  spine; 

no  ventral  fins;  skin  of  the  sides  is  very  hard Triggerfish,  p.  520 

First  dorsal  spine  is  not  much  stouter  than  the  others  and  cannot  be  locked  erect  by  the  second  spine;  ventral  fins 
are  well-developed;  skin  of  the  sides  is  soft 8 

8.  The  space  between  the  two  dorsal  fins  is  nearly  as  long  as  the  first  dorsal  fin,  or  longer;  the  ventral  fins  are  situated 

behind  the  middle  of  the  pectorals 9 

There  is  little  or  no  free  space  between  the  two  dorsal  fins;  the  ventrals  are  in  front  of  the  middle  of  the  pectorals 11 

9.  Jaws  long;  teeth  large  and  strong;  anal  with  one  spine Barracuda,  p.  306 

Jaws  short;  teeth  weak 10 

10.  Anal  fin  is  about  as  long  as  head  (snout  to  gill  openings)  and  has  one  weak  spine- refer  to  Silverside  family  key,  p.  302 
Anal  fin  is  only  about  half  as  long  as  head  and  has  three  stiff  spines  (only  two  spines  in  very  young  specimens). 

Mullet,  p.  305 

11.  Caudal  peduncle  is  extremely  slender;  the  caudal  fin  is  deeply  forked Pompano  family  (in  part)  key,  p.  371 

Caudal  peduncle  is  at  least  moderately  deep  and  thick;  the  caudal  fin  is  only  moderately  forked,  at  most 12 

12.  First  dorsal  fin  is  much  lower  than  second  dorsal 13 

First  dorsal  fin  is  as  high  as  the  second  dorsal  fin,  or  higher 14 

13.  Anal  fin  is  nearly  as  long  as  second  dorsal  fin Bluefish,  p.  383 

Anal  fin  is  only  about  one  half  as  long  as  second  dorsal  fin Rudderfish  (Pompano  family  in  part),  p.  373 

14.  Body  very  thin  through,  flat  sided,  nearly  two-thirds  as  deep  as  it  is  long  to  base  of  caudal  fin;  the  back  and  also  the 

ventral  edge  of  the  body  are  armed  with  bony  plates;  there  is  a  finlet  of  three  short  spines  in  front  of  the  anal 

fin.. John  Dory,  p.  297 

Body  stout,  not  more  than  one-third  as  deep  as  it  is  long;  the  sides  are  rounded;  the  back  and  lower  surface  are 
not  armed  with  bony  plates;  there  is  no  finlet  in  front  of  the  anal  fin 15 


8  FISHERY   BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 

Key  E — Continued 

15.  First  dorsal  fin  soft-rayed;  second  dorsal  fin  at  least  5  times  as  long  as  first  dorsal. 

refer  to  Cod  family  key,  in  part,  p.  173 
First  dorsal  fin  noticeably  spiny;  second  dorsal  fin  as  long  as  first  dorsal 16 

16.  Second  dorsal  fin  not  much  longer  than  the  anal  fin refer  to  Sea  bass  family  key,  in  part,  p.  389 

Second  dorsal  fin  is  about  twice  as  long  as  the  anal  fin refer  to  Weakfish  family  key,  p.  417 

Key  F 

Bony  fishes  with  snouts  of  ordinary  form;  symmetrical  tails;  caudal  fin  distinct  from  the  anal  fin;  neither  with  bar- 
bels on  the  top  of  the  snout  nor  with  canine  tusks;  and  with  only  one  well-developed  dorsal  fin;  the  latter  is  soft- 
rayed  except  that  it  may  be  preceded  by  a  few  short  spines  or  by  a  series  of  hair-like  rays  without  connecting 
membrane  and  that  there  may  be  an  isolated  spine  on  the  top  of  the  head  (from  No.  26,  p.  6). 

1.  The  rear  parts  of  the  dorsal  fin  and  of  the  anal  fin  are  broken  up  into  series  of  almost  separate  finlets  (fig.  191) 2 

The  rear  parts  of  the  dorsal  and  anal  fins  are  not  broken  up  into  series  of  finlets 3 

2.  The  forward  parts  of  the  dorsal  and  anal  fins  are  very  high  and  scythe-shaped;  the  pectorals  are  very  long,  reaching 

back  considerably  beyond  the  high  part  of  the  dorsal  fin;  there  are  no  spines  in  front  of  the  anal  fin. .Sea  bream,  p.  361 
The  dorsal  and  anal  fins  are  not  very  high  and  slope  gradually  rearward;  the  pectorals  are  small,  their  tips  falling 
far  short  of  the  level  of  the  front  of  the  dorsal  fin;  the  anal  fin  is  preceded  by  two  short  stout  spines. 

Leather  jacket,  p.  380 

3.  The  mouth  gapes  back  far  beyond  the  eye refer  to  Anchovies  key,  p.  118 

The  mouth  does  not  gape  back  much  beyond  the  rear  edges  of  the  eyes,  if  that  far 4 

4.  The  whole  of  the  anal  fin  is  behind  the  rear  end  of  the  dorsal  fin refer  to  Herring  Tribe  key,  p.  85 

Part  or  all  of  the  anal  fin  is  further  forward  than  the  rear  end  of  the  dorsal  fin 5 

5.  There  is  a  spine  or  a  bristle-like  rod  on  the  top  of  the  head  over  the  eyes 6 

There  is  no  spine  or  bristle-like  rod  on  the  head  over  the  eyes,  but  there  may  be  a  few  short  spines  close  in  front  of 

the  dorsal  fin 7 

6.  The  spine  on  the  top  of  the  head  is  thick  and  very  stiff  and  has  no  fleshy  tab  at  its  tip;  mouth  small;  body  stiff;  fin 

rays  slender,  not  fleshy refer  to  Filefish  family,  key,  p.  521 

The  spine  on  the  head  is  slender  and  flexible  and  has  a  fleshy  tab  or  "bait"  at  its  tip;  body  soft;  mouth  very  large; 
fin  rays  thick  and  fleshy Deep-sea  angler,  p.  543 

7.  Form  eel-like;  snout  sharp  pointed Launce,  p.  488 

Form  not  eel-like;  snout  blunt 8 

8.  Dorsal  fin  originates  on  the  head,  about  over  the  eyes Dolphin,  p.  360 

Dorsal  fin  originates  far  behind  the  eyes 9 

9.  Each  ventral  fin  is  represented  by  a  single  large  stout  spine refer  to  Stickleback  key,  p.  307 

The  ventral  fins  are  of  ordinary  rayed  type,  or  are  lacking 10 

10.  The  upper  anterior  profile  of  the  head  is  conspicuously  concave 11 

The  upper  anterior  profile  of  the  head  is  more  or  less  convex __12 

11.  The  forward  parts  of  the  dorsal  fin  and  of  the  anal  fin  are  much  higher  than  the  rear  parts,  the  first  few  rays  of  each 

being  very  much  longer  than  the  rays  farther  back Lookdown,  p.  379 

The  dorsal  and  anal  fins  are  only  a  little  higher  in  front  than  rearward,  the  first  few  rays  not  being  much  longer  than  the 
rays  farther  to  the  rear Moonfish,  p.  378 

12.  The  forward  rays  of  the  dorsal  and  anal  fins  are  very  long  and  thread-like Thread-fin,  p.  381 

The  forward  rays  of  the  dorsal  and  anal  fins  are  not  very  long  and  thread-like 13 

13.  The  entire  body  is  armored  with  several  rows  of  overlapping  plates Alligator  fish,  p.  457 

The  body  is  not  armored  with  overlapping  plates 14 

14.  The  skin  is  rough  or  prickly Refer  to  Puffers  and  Porcupine  fishes,  key,  p.  526 

The  skin  is  smooth,  though  scaly 15 

15.  The  front  part  of  the  dorsal  fin  is  much  higher  than  the  rear  part 16 

The  front  part  of  the  dorsal  fin  is  not  much  higher  than  the  rear  part 17 

16.  The  ventral  fins  are  large  and  conspicuous Opah,  p.  247 

There  are  no  ventral  fins Refer  to  Butterfish  and  Harvest  Fish,  key,  p.  363 

17.  The  tail  fin  is  conspicuously  rounded 18 

The  tail  fin  is  more  or  less  deeply  forked 19 

18.  The  dorsal  fin  runs  the  whole  length  of  the  back  from  close  behind  the  head  to  the  caudal  fin  which  it  joins;  there 

is  a  barbel  on  the  chin Cusk,  p.  238 

The  dorsal  fin  occupies  only  about  one-third  of  the  length  of  the  back  or  less,  and  stands  far  to  the  rear;  there  is  a 
considerable  space  between  it  and  the  caudal  fin;  there  is  no  barbel  on  the  chin. .Refer  to  Mummichog  key,  p.  162 


FISHES   OF  THE  GULF  OF   MAINE  9 

Key  F — Continued 

19.  The  caudal  peduncle  is  slender  and  has  a  conspicuous  longitudinal  keel  on  either  side;  the  pectoral  fins  do  not  reach 

back  as  far  as  the  point  of  origin  of  the  dorsal  fin Pilotfish,  p.  372 

The  caudal  peduncle  is  deep  and  has  no  longitudinal  keel;  the  pectoral  fins  reach  back  farther  than  the  point  of 
origin  of  the  dorsal  fin 20 

20.  There  are  6-8  short  detached  spines,  each  with  a  small  triangular  fin  membrane,  on  the  back  in  front  of  the  dorsal 

fin Barrelfish,  p.  369 

There  are  no  detached  spines  on  the  back  in  front  of  the  dorsal  fin 21 

21.  The  ventral  fins  stand  far  behind  the  bases  of  the  pectoral  fins;  the  point  of  origin  of  the  dorsal  fin  is  little  if  any  in 

advance  of  the  anal  fin;  the  pectoral  fins  (Gulf  of  Maine  species)  are  very  long,  reaching  back  nearly  to  the  base 

of  the  tail  fin Flying  fish,  p.  172 

The  ventral  fins  stand  about  under  the  base  of  the  pectoral  fins;  the  point  of  origin  of  the  dorsal  fin  is  far  in  ad- 
vance of  the  anal  fin;  the  pectoral  fins  are  small,  falling  far  short  of  the  anal  fin Black  ruff,  p.  370 

KeyG 

Fishes  as  in  Key  F,  except  that  at  least  the  forward  one-third  of  the  single  dorsal  fin  is  spiny.     There  is  no  adipose  fin 
behind  the  rayed  dorsal  nor  fleshy  flap  in  front  of  it  (from  No.  26,  p.  6). 

1.  The  body  (tip  of  snout  to  base  of  caudal  fin)  is  at  least  as  deep  as  it  is  long Boarfish,  p.  438 

The  body  is  considerably  longer  than  it  is  deep 2 

2.  The  rear  part  of  the  dorsal  fin  is  soft-rayed 3 

The  whole  length  of  the  dorsal  fin  is  spiny 8 

3.  Sides  of  head  bony,  with  knobs  or  spines 4 

No  knobs  or  spines  on  the  sides  of  the  head 5 

4.  Sides  of  head  armed  with  conical  spines;  the  spiny  portion  of  the  dorsal  fin  is  at  least  as  long  as  the  soft  part;  the 

body  is  flattened  sidewise Refer  to  Rosefish  family,  key,  p.  430 

Sides  of  head  with  low  rounded  knobs  only;  the  spiny  portion  of  the  dorsal  fin  is  considerably  shorter  than  the  soft 
part;  body  tadpole-shaped Arctic  sculpin  (Sculpin  family  in  part),  p.  453 

5.  The  ventral  fins  are  much  larger  than  the  pectorals;  the  eyes  are  very  large Short  big-eye,  p.  410 

The  ventral  fins  are  not  larger  than  the  pectorals ;  the  eyes  are  not  very  large 6 

6.  The  pectorals  are  sharply  pointed;  the  body  is  much  flattened  sidewise Refer  to  Porgy  family,  key,  p.  411 

The  pectorals  are  rounded;  the  body  is  not  much  flattened  sidewise 7 

7.  The  rear  (soft)  part  of  the  dorsal  fin  is  nearly  as  long  as  the  front  (spiny)  part;  the  anal  fin  is  much  higher  than 

long Seabass  (Seabass  family  in  part),  p.  407 

The  rear  (soft)  part  of  the  dorsal  fin  is  less  than  half  as  long  as  the  spiny  (front)  part;  the  anal  fin  is  longer  than 
high Refer  to  Cunner  family,  key,  p.  473 

8.  The  mouth  is  strongly  oblique;  there  are  no  ventral  fins Wrymouth,  p.  500 

The  mouth  is  not  strongly  oblique;  ventral  fins  are  present  (very  small  in  one  species).. Refer  to  Blenny  fishes,  key,  p.  491 

THE  CYCLOSTOMES.    CLASS  AGNATHA 

The  lampreys  are  the  most  primitive  of  the  appearance,  but  are  easily  distinguishable  from 

true  vertebrates,  their  skeletons  being  cartilagi-  the  true  eels  and,  indeed,  from  most  of  the  true 

nous   without   any   true  bone,   and   their  skulls  fishes,  by  their  peculiar  jawless  sucking  mouth 

hardly  differentiated  from  the  vertebral  column  situated  at  the  tip  of  the  snout,  and,  further, 

which  forms  a  simple  notochordal  sheath.    They  from  all  Gulf  of  Maine  eels  by  lacking  pectoral 

have  no  true  jaws,  no  ribs,  no  shoulder  or  pelvic  fins, 
girdles,  and  no  paired  fins.     They  are  eel-like  in 

THE  HAGFISHES  AND  LAMPREYS.    FAMILIES  MYXINIDAE  AND  PETROMYZONIDAE 

These  two  groups  are  easily  distinguished,  one  nally,  whereas  the  lampreys  have  no  barbels,  their 

from  the  other,  by  the  fact  that  the  hags  have  mouths  are  disc-  or  funnel-like,  their  eyes  are  well 

several  barbels  on  the  chin,  that  their  mouths  developed  after  the  larval  stage  is  past,  and  they 

are  not  disc-  or  funnel-like,  that  they  have  only  have  one  or  more  dorsal  fins  separate  from  the 

one  continuous  fin  fold  on  the  back  and  around  caudal  fin. 
the  tail,  and  that  their  eyes  are  not  visible  exter- 


10 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Hagfish  Myxine  glutinosa  Linnaeus  1758 
Bigelow  and  Schroeder,  1948,  p.  34. 

Description. — The  hag,  like  the  lamprey,  lacks 
paired  fins  and  fin  rays.  Its  skeleton  is  wholly 
cartilaginous,  without  bones,  its  mouth  is  jawless; 
and  its  skin  is  scaleless.  It  is  easily  recognized 
by  its  eel-like  form;  by  its  single  finfold  (a  fold 
of  skin,  not  a  true  fin)  running  right  around  the 
tail  and  forward  on  the  lower  surface  of  the  body 
with  no  division  into  dorsal,  caudal,  and  anal  fins; 
by  the  single  gill  pore  on  each  side,  just  forward 
of  the  origin  of  the  ventral  finfold;  by  its  lipless 
mouth,  star-shaped  in  outline  when  closed;  by 
the  single  nasal  aperture  at  the  tip  of  the  snout; 
by  its  peculiar  barbels  or  "tentacles,"  two  flanking 
the  mouth  on  either  side  and  four  surrounding 
the  nostril;  and  by  the  evertible  tongue  studded 
with  rows  of  horny  rasplike  "teeth."  We  might 
also  mention  the  series  of  mucous  sacs  on  either 
side  of  the  abdomen,  and  point  out  that  the  dorsal 
finfold  originates  about  two-thirds  of  the  distance 
back  from  snout  toward  tip  of  tail,  and  the  ven- 


tral fin  fold  one-third  the  way  back,  with  the  vent 
piercing  it. 

Color. — Hags  vary  in  color,  perhaps  to  cor- 
respond with  the  color  of  the  bottom,  being  gray- 
ish brown  or  reddish  gray  above,  variously  suf- 
fused, mottled,  or  piebald  with  darker  or  paler 
gray,  with  brown,  or  with  bluish;  they  are  whitish 
or  pale  gray  below. 

Size. — Gulf  of  Maine  hags  grow  commonly  to  a 
length  of  about  1%  to  2  feet,  with  a  maximum  of 
31  inches  recorded  off  the  coast  of  Maine. 

Habits. — The  hag  is  found  chiefly  if  not  ex- 
clusively where  the  bottom  is  soft  mud,  where  (to 
judge  from  its  actions  during  the  brief  time  it 
survives  in  aquaria)  it  spends  its  time  lying  em- 
bedded in  the  clay  or  mud  with  the  tip  of  the 
snout  projecting.  And  it  is  at  home  only  in  com- 
paratively low  temperatures,  cooler  probably, 
than  50°,  which  confines  it  in  summer  to  depths 
of  15  to  20  fathoms  or  more  in  the  Gulf  of  Maine. 
It  is  not  a  true  parasite,  as  has  sometimes  been 
suggested,  their  being  no  reason  to  believe  it  ever 
attacks  living,  uninjured  fish,  but  is  a  scavenger. 


Figure  2. — Hagfish  (Myxine  glutinosa).  A,  adult,  Gulf  of  Maine,  from  Bigelow  and  Schroeder,  drawing  by  E.  N.  Fischer. 
B,  lower  view  of  head  of  same;  C,  tongue-teeth  of  same  as  seen  from  above,  about  3  times  natural  size;  D,  egg, 
after  Dean,  about  2  times  natural  size. 


FISHES   OF  THE  GULF  OF   MAINE 


11 


Being  blind,  it  doubtless  finds  its  food  by  its 
greatly  specialized  olfactory  apparatus.  It  feeds 
chiefly  on  fish,  dead  or  disabled,  though  no  doubt 
any  other  carrion  would  serve  it  equally  well. 
And  it  is  known  to  prey  on  marine  annelid  worms 
also,  at  least  in  Norwegian  waters.  It  is  best 
known  for  its  troublesome  habit  of  boring  into 
the  body  cavities  of  hooked  or  gilled  fishes,  eat- 
ing out  the  intestines  first  and  then  the  meat,  and 
leaving  nothing  but  a  bag  of  skin  and  bones,  inside 
of  which  the  hag  itself  is  often  hauled  aboard,  or 
clinging  to  the  sides  of  a  fish  it  has  just  attacked. 
In  fact,  it  is  only  in  this  way,  or  entangled  on 
lines,  that  hags  ordinarily  are  taken  or  seen. 

Being  worthless  itself,  the  hag  is  an  unmitigated 
nuisance,  and  a  particularly  loathsome  one  owing 
to  its  habit  of  pouring  out  slime  from  its  mucous 
sacs  in  quantity  out  of  all  proportion  to  its  small 
size.  One  hag,  it  is  said,  can  easily  fill  a  2-gallon 
bucket,  nor  do  we  think  this  any  exaggeration. 

In  American  waters  the  commercial  fishes  most 
often  damaged  by  it  are  haddock  and  the  hakes 
(Urophycis),  these  being  the  species  most  often 
fished  for  with  long  lines  or  with  gill  nets  over  the 
type  of  bottom  the  hag  frequents.  But  it  some- 
times damages  cod  also,  and  European  authors 
describe  it  as  attacking  ling  (Molva)  and  other 
members  of  the  cod  tribe,  herring,  mackerel, 
sturgeon,  and  even  mackerel  sharks  under  similar 
circumstances. 

Breeding  habits.— The  hag  was  formerly  believed 
to  be  a  functional  hermaphrodite,  with  its  single  sex 
organ  first  developing  sperm  in  the  posterior  por- 
tion, eggs  later  in  the  anterior  portion.  However, 
recent  detailed  studies  of  the  sex  organ  appear  to 
show  that  such  is  not  the  case,  but  that  either  the 
male  portion  of  the  common  sex  organ  matures 
in  a  given  individual  with  the  female  portion 
remaining  rudimentary,  or  vice  versa.13 

It  has  long  been  known  that  the  eggs  are  few 
in  number  (only  19  to  30  having  been  counted  in 
any  one  female)  and  large  (up  to  25  mm.  in  length), 
and  the  horny  shell  has  a  cluster  of  anchor-tipped 
filaments  at  each  end  that  make  the  eggs  easy  of 
identification.  Until  1900  none  had  been  found 
that  certainly  had  been  laid  naturally.  In  that 
year,  however,  hag  eggs  were  reported  from  the 
western  part  of  Georges  Bank  and  from  the  south 

u  See  Blgelow  and  Sihroeder,  Fishes  Western  North  Atlantic,  Pt.  1,  ch.  2, 
1948,  pp.  35-36,  for  references. 


coast  of  Newfoundland  by  Dean  (1900) ;  M  from  the 
neighborhood  of  the  Faroe  Islands  by  Jensen; 1S 
from  Norway  by  Hjort;  16  off  Morocco  bv  Koe- 
foed.17  And  they  have  been  reported  subsequently 
from  the  Bay  of  Fundy  by  Huntsman,  from 
Frenchman  Bay  on  the  coast  of  Maine  by  Conel.18 
The  eggs  are  deposited  on  bottom,  where  they  stick 
firmly  to  fixed  objects  of  one  sort  or  another  by 
their  terminal  filaments  and  by  threads  of  slime. 

The  hag  spawns  throughout  its  range;  also  it 
spawns  throughout  the  year,  for  females  nearing 
ripeness  and  others  nearly  spent  have  been  re- 
corded for  winter  and  spring,  as  well  as  summer 
and  autumn,  in  one  part  of  its  range  or  another. 
The  few  eggs  so  far  reported  have  been  from  depths 
of  50  to  150  fathoms,  most  of  them  trawled  on 
mud,  clay,  or  sand  bottom. 

We  need  only  add  that,  to  judge  from  their 
behavior  in  aquaria,  the  females  cease  to  feed  at 
the  approach  of  sexual  maturity,  as  many  other 
fishes  do.  Newly  hatched  hags  have  never  been 
seen,  but  inasmuch  as  the  smallest  yet  described 
(about  2%  inches  long),  probably  not  long  out  of 
the  egg,  already  resembled  the  adult  in  external 
appearance  there  is  no  reason  to  suppose  that  the 
hag  passes  through  a  larval  stage  greatly  different 
from  the  adult. 

General  range. — Arctic  seas,  and  both  coasts  of 
the  north  Atlantic;  Murman  Coast  and  northern 
Norway  south  regularly  to  the  Irish  Sea,  and  to 
Morocco  as  a  stray  in  the  East;  northern  part  of 
Davis  Strait,  south  to  the  latitude  of  Cape  Fear, 
N.  C,  in  the  west.  It  is  represented  in  the  cor- 
responding temperature-belt  of  the  Southern 
Hemisphere  by  a  form  (or  forms)  resembling  it  so 
closely  that  it  is  doubtful  whether  any  sharp  line 
can  be  drawn  between  them. 

Occurrence  in  the  Gulf  of  Maine. — Apart  from 
one  record  for  the  northern  part  of  Davis  Strait, 
the  most  northerly  reports  of  the  hag  off  the 
American  coast  are  from  southern  Newfoundland 
and  from  the  Grand  Banks.19  But  it  is  generally 
distributed  along  outer  Nova  Scotia  at  appropriate 
depths.     And  it  is  only  too  common  in  the  Gulf 

»  Mem.  N.  Y.  Acad.  Set.,  vol.  2,  Pt.  2,  Art.  2,  1900. 

"  Vlden.  Meddel.  Dansk  naturhlst.    Forenlng,  1900,  p.  1. 

"  Rept.  Norwelglan  Fishery  and  Mar.  Invest.,  vol.  1, 1900,  No.  1,  ch.  4,  p.  75. 

«  Rept.  Michael  Sars  North  Atlantic  Exped.,  Zool.,  vol.  4,  No.  1, 1927,  p.  18 

"  Science,  N.  Ser.,  vol.  75, 1932,  pp.  19-20. 

<•  It  has  not  been  reported  for  certain  from  West  Greenland  (so  far  as  we 
can  learn),  from  the  outer  coast  of  Labrador,  or  within  the  Gulf  of  St. 
Lawrence  though  It  Is  to  be  eipected  in  the  deeper  parts  of  the  latter. 


12 


FISHERY  BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


of  Maine;  perhaps  it  is  not  absent  there  from 
any  considerable  area  of  smooth  bottom.  Thus, 
it  is  abundant  off  the  north  end  c  f  Grand  Manan ; 
is  reported  from  Passaamquoddy  Bay  and  from 
various  localities  near  Eastport;  is  to  be  found 
off-shore  on  muddy  bottom  all  along  the  Maine 
coast;  and  is  caught  at  times  in  considerable 
numbers  on  the  Boon  Island-Isles  of  Shoals 
fishing  grounds  and  about  Jeffreys  Ledge,  where 
we  found  it  plentiful  enough  in  the  spring  of  1913 
to  have  gutted  3  to  5  percent  of  all  the  haddock 
in  the  gill  nets.  Fishermen  report  it  as  equally 
numerous  in  the  deeper  parts  of  Massachusetts 
Bay.  On  the  offshore  banks  the  hag  is  well  known, 
and  it  has  been  trawled  at  various  localities 
along  the  outer  edge  of  the  Continental  Shelf  off 
New  England  at  depths  of  from  100  to  200 
fathoms,  and  deeper.  We  ourselves  took  11 
large  ones  in  one  set  of  a  Monaco  deep-sea  trap 
in  260  fathoms  off  Nantucket  on  July  9,  1908, 
and  it  has  been  taken  in  from  300  to  500  fathoms 
off  Marthas  Vineyard;  as  deep  as  524  fathoms  on 
the  southeast  slope  of  Georges   Bank. 

Sea  lamprey  Petromyzon  marinus  Linnaeus  1758 

Lamprey;    Spotted    lamprey;    Lamper;    Eel- 
sucker 

Bigelow  and  Schroeder,  1948,  p.  46. 

Description. — Lampreys  are  eel-like  in  ap- 
pearance, but  have  a  soft,  cartilaginous  skeleton. 
They  lack  paired  fins  but  have  well  developed 


dorsal  and  ventral  finfolds.  In  the  adult  the 
jaws  are  so  rudimentary  that  apparently  they 
are  wanting;  the  mouth  is  a  longitudinal  slit 
when  closed,  but  forms  an  elliptical  disk  at  the 
tip  of  the  snout  when  open,  and  is  armed  with 
many  horny,  hooked  teeth  arranged  in  numerous 
(11  to  12)  rows,  the  innermost  the  largest.  There 
are  two  dorsal  finfolds,  and  there  are  seven  open 
gill  slits  on  each  side,  whereas  the  hag  has  only 
one  gill  pore  on  each  side,  and  only  one  fin.  The 
sea  lamprey  (the  only  member  of  its  group  known 
from  our  salt  waters)  can  hardly  be  mistaken 
for  any  other  fish,  its  eel-like  appearance  coupled 
with  two  dorsal  fins  and  the  jawless  mouth 
placing  it  at  a  glance. 

Color. — Small  specimens  (whether  on  their 
way  downstream  or  in  salt  water)  are  white  below 
and  uniformly  colored  above,  usually  described 
as  blackish  blue,  or  as  lead  colored,  and  more 
or  less  silvery.  But  large  specimens  usually  are 
olive  brown  above,  or  of  varying  shades  of  yellow- 
brown,  green,  red,  or  blue,  mottled  with  a  darker 
shade  of  the  same  color,  or  sometimes  nearly 
black  if  the  dark  patches  are  confluent.  The 
lower  surface  is  whitish,  gray,  or  of  a  pale  shade 
of  the  same  hue  as  the  ground  color  of  the  back. 
During  breeding  season,  the  landlocked  form 
takes  on  more  brilliant  hues,  with  the  ground  tint 
turning  bright  yellow. 

Size. — The  length  at  the  time  of  transformation 
from  the  larval  stage  is  about  4  to  8  inches  (100- 
200  mm.).     Sexually  mature  individuals,   taken 


'* 


"^M^** 


Figure  3. — Sea  lamprey  {Petromyzon  marinus),  about  18  inches  long,  Merrimac  River;  and  open  mouth  disc  of  another 
Merrimac  River  specimen  to  show  the  arrangement  of  the  horny  teeth,  about  0.9  times  natural  size.  From  Bigelow 
and  Schroeder.     Drawings  by  E.  N.  Fischer. 


FISHES   OF  THE  GULF  OF   MAINE 


13 


in  American  rivers,  average  2  to  2%  feet  long, 
up  to  a  maximum  of  about  3  feet.  One  of  33 
inches  weighed  2%  pounds. 

Habits. — It  has  been  known  from  early  times 
that  the  sea  lamprey  breeds  in  fresh  water.  How- 
ever, it  does  not  enter  all  the  streams  within  its 
range  indiscriminately.  As  an  illustration,  we 
may  cite  outer  Nova  Scotia  and  the  Bay  of  Fundy, 
where  lampreys  run  in  the  St.  Marys,  Sackville, 
Annapolis,  Shubenacadie,  Petit  Codiac,  and  St. 
Johns  Rivers,  but  not  in  the  Moser  or  Apple 
Eivers,  although  these  last  also  are  "salmon" 
rivers.  Their  requirements  are  a  gravelly  bottom 
in  rapid  water  for  their  spawning  beds,  with  muddy 
or  sandy  bottom  in  quiet  water  nearby,  for  the 
larvae. 

In  many  small  streams,  and  in  larger  ones  if 
these  are  blocked  by  dams  or  high  falls,  they  may 
spawn  only  a  short  distance  upstream ;  even  within 
the  influence  of  the  tide,  although  invariably  in 
fresh  water.  But  they  are  able  to  ascend  falls, 
if  these  are  not  too  steep  and  high,  by  clinging 
to  the  rocks  by  their  oral  discs  and  resting.  And 
they  may  run  upstream  for  very  long  distances  in 
large  rivers,  as  they  did  formerly  in  the  Merrimac 
and  probably  still  do  in  the  St.  Johns  River.  They 
are  still  to  be  found  200  miles  or  more  from  the  sea 
in  the  upper  tributaries  of  the  Delaware  and  Sus- 
quehanna systems. 

Since  the  breeding  activities  of  the  sea  lamprey 
take  place  in  fresh  water,  a  brief  account  will 
suffice  here.  As  the  two  sexes  ripen,  the  males 
develop  a  strong  ridge  along  the  back,  the  females 
a  crestlike  fin  between  the  anus  and  the  caudal 
fin.  Spawning,  commencing  when  the  tempera- 
ture of  the  water  is  about  50°  F.  (10°  C.)  is  com- 
pleted by  the  time  it  has  warmed  to  about  68°-70° 
(20°-21°  C),  and  a  sea  lamprey  has  been  found 
to  contain  236,000  ova.  Working  in  pairs,  some- 
times with  a  second  female  assisting,  they  make 
depressions  2  to  3  feet  in  diameter  and  about  6 
inches  deep  in  the  stream  bed  in  stretches  where 
the  bottom  is  stony  or  pebbly,  dragging  the  stones 
downstream  in  a  pile  with  their  suckerlike 
mouths.  And  they  are  able  to  move  stones  as 
large  as  one's  fist.  It  is  in  these  depressions  that 
the  eggs  are  deposited,  not  among  the  piles  of 
discarded  stones  that  have  often  been  described 
as  "nests."  It  seems  that  they  all  die  after  spawn- 
ing; not  only  have  they  often  been  found  dead, 
but  their  intestines  atrophy,  they  are  attacked  by 


fungus,    and    they    become    so    debilitated    that 
recovery  seems  out  of  the  question. 

The  larvae  are  different  in  appearance  from  the 
adults:  blind,  toothless,  with  mouths  and  fins  of 
different  shape.  They  continue  in  this  state  for 
a  period  estimated  as  3  to  4  years,  during  most  of 
which  time  they  live  in  burrows  in  the  mud  or 
sand,  or  hide  under  stones.  They  are  abundant 
in  the  mud  of  flats  near  the  mouths  of  small 
tributary  streams  of  river  systems  such  as  the 
Delaware  and  Susquehanna,  where  lampreys  still 
breed  in  large  numbers,  and  they  subsist  on 
minute  organisms.  At  the  end  of  this  larval  period, 
when  they  have  grown  to  a  length  of  4  to  6  inches, 
they  undergo  transformation  to  the  adult  form 
and  structure,  an  event  occupying  about  two 
months,  August  to  September  or  October.  They 
run  down  to  the  sea  in  November  or  December,  to 
live  and  grow  there  for  one  or  two  years,  so  that 
large  ones,  not  yet  mature,  are  to  be  found  in 
salt  water  all  the  year  round. 

Little  is  known  of  the  habits  of  the  lampreys 
while  they  live  in  the  sea  further  than  that  their 
mode  of  life  centers  around  a  fiercely  predaceous 
nature.  Judging  from  their  land-locked  relatives 
and  from  the  occasions  on  which  they  have  been 
found  fastened  to  sea  fish,  they  must  be  extremely 
destructive  to  the  latter,  which  they  attack  by 
"sucking  on"  with  their  wonderfully  effective 
mouths.  The  lamprey  usually  fastens  to  the  side 
of  its  victim,  where  it  rasps  away  until  it  tears 
through  the  skin  or  scales  and  is  able  to  suck  the 
blood.  Its  prey  sucked  dry,  it  abandons  it  for 
another.  Probably  lampreys  are  parasites  and 
bloodsuckers  pure  and  simple,  for  we  cannot  learn 
that  anything  but  blood  has  been  found  in  their 
stomachs,  except  fish  eggs,  of  which  lampreys  are 
occasionally  full.20 

In  salt  water  they  have  been  found  preying  on 
mackerel,  the  various  anadromous  herrings,  cod, 
haddock,  American  pollock  (Pollachius) ,  salmon, 
basking  sharks,  swordfish,  hake  (Urophycis), 
sturgeons  and  eels.  Sometimes  as  many  as  three 
or  four  are  fast  at  one  time  to  a  single  shad,  and 
they  are  said  to  be  exceedingly  aggressive  in 
their  attacks  on  other  fishes.  Occasionally  they 
are  found  fast  to  driftwood,  even  to  boats.  When 
not  clinging  to  anything  they  are  strong,  vigorous 
swimmers,  progressing  by  an  undulating  motion. 


»  Ooode,  Fish.  Ind.  U.  S.,  Sect.  1, 1884,  p.  677. 


14 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


General  range. — Atlantic  coasts  of  Europe  and  of 
North  America;  from  the  west  coast  of  Greenland 
to  Florida  in  the  western  side  of  the  Atlantic ;  from 
northern  Norway  to  the  Mediterranean  in  the 
eastern;21  running  up  fresh  rivers  to  breed,  and 
landlocked  in  certain  American  lakes. 

Occurrence  in  the  Gulf  of  Maine. — No  doubt  the 
sea  lamprey  occurs  along  the  whole  coast  line  of  the 
Gulf  of  Maine,  for  it  is  recorded  in  or  at  the  mouths 
of  numerous  rivers  and  streams  in  Nova  Scotia, 
New  Brunswick,  Maine,  and  Massachusetts;  spe- 
cifically in  the  St  John,  Annapolis,  PetitCodiac,  and 
Shubenacadie  Rivers  and  from  the  St.  Andrews 
region  in  salt  water  in  the  Bay  of  Fundy;  from 
Eastport,  Bucksport,  Casco  Bay,  and  the  Pre- 
sumpscott,  Kennebec,  and  Penobscot  Rivers  in 
Maine;  from  the  Merrimac  River  system;  from  the 
Exeter  and  Lamprey  Rivers,  tributaries  of  Great 
Bay,  New  Hampshire ;  and  from  the  Parker  River 
in  northern  Massachusetts. 

Since  lampreys  never  take  the  hook  or  are  cap- 
tured in  nets  except  on  rare  occasions  they  are  sel- 
dom seen  in  salt  water;  only  when  running  up  our 
rivers  are  they  familiar  objects.  But  they  have 
been  taken  as  far  offshore  as  tho  seaward  slopes  of 
Banquereau,  Sable  Island,  and  LaHave  Banks  off 
Nova  Scotia;  on  Browns  Bank;  in  the  deep  gully 
between  the  latter  and  Georges  Bank,  and  over  the 
continental  slope  off  Nantucket  and  off  Marthas 
Vineyard. 

Lampreys  have  long  been  known  to  run  up  New 
England  rivers  a  little  earlier  in  the  spring  than 
shad,  perhaps  beginning  to  work  upstream  as  early 
as  the  beginning  of  April  or  even  the  end  of  March. 
In  the  rivers  tributary  to  the  Gulf  of  Maine  the 
runs  are  at  their  peak  during  May  and  early  June, 
with  few,  if  any,  entering  later  than  that.  The 
larvae  have  been  reported  by  Doctor  Huntsman 
as  plentiful  in  the  Shubenacadie  (emptying  into 
the  Bay  of  Fundy)  and  no  doubt  they  are  to  be 
found  in  the  Merrimac  system,  in  the  Exeter  River, 
and  in  other  Gulf  of  Maine  streams. 

Abundance. — The  construction  of  impassible 
dams  has  sadly  reduced  the  numbers  of  lampreys 

si  Also  reported  from  "West  Africa"  by  Oflnther,  Cat.  Fishes  British  Mu- 
seum, vol.  8, 1870,  p.  502. 


in  the  larger  rivers  of  New  England.  In  the  Mer- 
rimac, for  example,  once  a  famous  lamprey  river,22 
so  few  now  succeed  in  surmouating  the  succession 
of  dams  that  a  recent  survey  yielded  no  evidence 
of  any  now  having  access  to  the  upper  reaches. 
Some  lampreys,  however,  are  said  to  breed  in  the 
river  below  the  Lowell  dam;23  we  have  seen  what 
resembled  their  "nests"  in  the  Squannacook,  a 
branch  of  the  Nashua  tributary  to  the  Middle  Mer- 
rimac, and  they  still  continue  numerous  in  some 
Gulf  of  Maine  streams  where  they  can  reach  suit- 
able spawning  grounds  without  too  great  difficulty. 
We  may  quote  catches  of  up  to  119  recently  in  the 
Shubenacadie,  where  larvae  also  have  recently 
been  reported  in  abundance,24  and  of  more  than 
100  each  on  several  occasions  in  the  Exeter 
River,25  where  they  are  familiar  spectacles,  as  they 
gather  at  the  falls  at  Exeter,  N.  H.  But  we  ought 
perhaps  to  caution  the  reader  that  while  lampreys, 
like  other  anadromous  fishes,  may  seem  plentiful 
when  condensed  between  the  narrow  bounds  of  a 
river's  banks,  their  numbers  as  a  whole  do  not  rival 
those  of  the  more  abundant  of  the  salt-water  fishes. 
Importance. — Lampreys  were  esteemed  a  great 
delicacy  in  Europe  during  the  middle  ages  (histo- 
rians tell  us  Henry  I  of  England  died  of  a  surfeit  of 
them)  and  considerable  numbers  were  captured  of 
old  in  the  rivers  of  New  England  for  human  food, 
particularly  in  the  Connecticut  and  Merrimac 
Rivers.  But  the  lamprey  fishery  has  been  scarcely 
more  than  a  memory  for  40  years  past  except  lo- 
cally and  in  a  small  way  for  home  consumption,  or 
to  supply  the  needs  of  biological  laboratories.  In 
the  salt  water  of  the  Gulf  of  Maine  the  lamprey  has 
never  been  of  any  commercial  importance;  the 
average  fisherman  might  not  see  one  in  a  lifetime, 
nor  is  there  any  sale  for  the  few  that  are  picked  up 
by  chance.  But  larvae  are  taken  in  considerable 
numbers  for  bait  in  the  Susquehanna  River,  and 
perhaps  elsewhere  along  the  middle  Atlantic  coast. 


aa  For  an  account  of  the  lamprey  fishery  in  New  England  during  the  first 
half  of  the  19th  century,  see  Qoode,  Fish,  and  Fishery  Ind.  U.  S.,  Sect.  1, 1884, 
p.  680. 

"  Bailey,  Biol.  Survey  Merrimack  Watershed,  New  Hampshire  Fish  and 
Game  Dept.,  1938,  p.  158. 

a*  Information  gathered  for  us  by  Dr.  A.  G.  Huntsman. 

a'  Collected  for  the  Biological  Laboratory,  Harvard  University. 


FISHES   OF  THE  GULF  OF   MAINE 


15 


CARTILAGINOUS    FISHES.      CLASS  CHONDRICHTHYES 

The  Shark  and  Skate  Tribes,  and  the  Chimaeroids 


These  are  fishlike  vertebrates  with  well-devel- 
oped fins  and  teeth,  and  with  2  pairs  of  fins,  one  of 
them  supported  by  the  pectoral  girdle,  the  other 
by  the  pelvic  girdle.  Their  most  distinctive  char- 
acter, as  contrasted  with  the  bony  fishes  (p.  80) 
is  that  their  entire  skeleton,  including  the  skull, 
is  cartilaginous,  without  any  true  bone,  though  it 
is  partly  calcified,  especially  in  the  vertebrae; 
the  skull  is  far  simpler  than  it  is  among  the  bony 
fishes;  the  gill  filaments  are  attached  throughout 
their  lengths  to  the  partitions  between  the  gill 
openings  instead  of  being  free;  and  the  rear  portion 


of  the  digestive  tract  is  modified  into  the  so-called 
"spiral  valve"  by  the  development  of  a  special  fold 
from  its  lining  layer,  which  only  a  few  bony  fishes 
have. 

Fertilization  is  internal  in  all  of  them,  and  is 
effected  by  a  pair  of  rodlike  copulatory  organs,  each 
of  which  is  developed  from  the  inner  edge  of  one  of 
the  two  pelvic  fins,  and  is  supported  by  one  or  more 
cartilages. 

The  sharks  and  rays  are  usually  looked  upon  as 
more  primitive  than  the  bony  fishes. 


SHARKS,  TORPEDOES,  SKATES,  AND  RAYS.     SUBCLASS  ELASMOBRANCHII 


The  most  obvious  external  character  by  which 
all  the  sharks,  skates,  and  rays  are  distinguishable 
from  all  of  the  bony  fishes  is  that  tbey  have  five  or 
more  gill  openings  on  either  side  of  the  head,  in- 
stead of  only  one.  They  recall  the  lampreys  in 
this  respect,  but  it  is  a  commonplace  that  their 
jaws  and  teeth  are  extremely  well-developed. 
Their  skins  are  tough,  and  are  studded  in  most  of 
them  with  denticles  (placoid  scales),  which  are  not 
homologous  with  the  scales  of  bony  fishes,  for  both 
dermis  and  epidermis  take  part  in  their  formation, 
instead  of  the  dermis  alone.  The  teeth  of  the 
sharks  and  rays  represent  placoid  scales  that  are 
modified  and  are  embedded  in  the  gums  alone,  not 
in  the  jaws.  The  fins  are  supported  at  their  bases 
by  segmented  cartilaginous  rods,  supplemented  in 
all  of  the  sharks,  and  in  some  of  the  rays  by  nu- 
merous slender  horny  fibers  further  out,  instead  of 
by  rays  or  spines  of  the  sorts  that  are  seen  in  the 
bony  fishes.  All  of  their  fins  are  covered  with  the 
same  leathery  skin  that  clothes  the  body.  Among 
sharks  the  tail  is  uneven  ( "he tero cereal"),  with 


the  vertebral  column  extending  out  into  its  upper 
lobe,  but  it  is  whip-like  in  most  of  the  skates  and 
rays,  with  no  definite  caudal  fin.  The  torpedo 
is  an  exception  to  this  rule. 

The  modern  representatives  of  the  subclass  may 
be  grouped  in  two  orders,  the  one  (Selachii)  to 
include  all  living  sharks,  the  other  (Batoidei)  to 
include  the  sawfishes,  the  skates  and  the  rays. 
They  are  separated  one  from  the  other  by  the 
following  external  differences,  and  there  are  skele- 
tal differences  between  them  as  well:26 

1.  The  gill  openings  are  at  least  partly  on  the  sides;  the 
edges  of  the  pectoral  fins  are  not  attached  to  the  sides  of  the 
head  in  front  of  the  gill  openings;  the  upper  edges  of  the 
orbits  are  free  from  the  eyeballs,  so  that  they  form  free 

eyelids Sharks,    (p.   15). 

The  gill  openings  are  entirely  on  the  lower  surface;  the 
edges  of  the  pectoral  fins  are  attached  to  the  side  of  the 
head  in  front  of  the  gill  openings;  the  upper  edges  of  the 
orbits  are  attached  to  the  eyeballs  so  that  they  do  not  form 
free   eyelids Sawfishes,  skates  and   rays,   (p.  57). 


*■  For  further  discussion,  see  Bigelow  and    Schroeder,  Fishes   Western 
North  Atlantic,  Ft.  1,  ch.  3,  1948,  p.  64. 


Sharks.     Order  Selachii 


Sharks  always  are  objects  of  interest,  not  only 
to  fishermen  and  mariners  but  to  seaside  visitors 
generally,  because  of  their  evil  appearance,  their 
ferocity,  the  large  size  to  which  some  of  them  grow, 
the  destruction  they  wreak  on  fishermen's  nets 
and  lines  as  well  as  on  the  smaller  fishes  on  which 
they  prey,  and  because  of  the  bad  reputation 
certain  kinds  have  earned  as  maneaters. 


The  Gulf  of  Maine  is  not  particularly  rich  in 
sharks  (very  poor  indeed  compared  with  our 
southern  coasts),  for  while  the  number  of  species 
actually  recorded  there  is  considerable  (indeed 
any  high-seas  shark  might  stray  thither)  the  little 
spiny  dogfish  alone  is  numerous  in  the  sense  in 
which  this  term  is  applied  to  the  various  com- 
mercial fishes.     And  only  two  of  the  larger  species, 


16 


FISHERY  BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


the  mackerel  shark  (Larrma  nasus),  and  the  blue 
shark  (Prionace  glauca),  occur  with  us  in  numbers 
sufficient  for  one  to  be  fairly  sure  of  seeing  them 
during  a  summer's  boating  off  the  coast  north  of 
Cape  Cod. 

With  the  larger  sharks  generally  so  scarce  (the 
mackerel  shark  is  harmless  to  anything  larger  than 
the  fishes  on  which  it  feeds,  and  the  blue 
shark  is  also  harmless,  although  better  armed), 
the  danger  of  attacks  on  bathers  is  negligible  in 
our  Gulf.  Indeed,  not  a  single  well-authenticated 
instance  of  the  sort  is  on  record  w  for  the  past 
80  years  for  the  coast  north  of  Cape  Cod,  though 
the  beaches  are  crowded  every  summer  with 
vacationists.  But  as  long  as  the  white  shark  or 
man-eater  (Carcharodon  carcharias)  does  stray 
occasionally  into  the  Gulf  (p.  26),  it  is  always 
remotely  possible  that  we  may  be  horrified  some 
summer  by  the  news  of  tragedies  such  as  occurred 
on  the  New  Jersey  coast  in  July  1916,  when 
several  persons  were  killed  or  injured,  presumably 
by  a  small  shark  of  this  species  that  was  captured 
nearby  a  few  days  later,28  and  near  Mattapoisett, 
on  Buzzards  Bav,  Mass.,  on  July  25,  1936,  when 
a  swimmer  was  fatally  injured  by  a  shark,  species 
not  determined.29 

17  In  1830  (an  event  often  quoted)  one  Joseph  Blaney,  fishing  from  a  small 
boat  In  Massachusetts  Bay  off  Swampscott,  Mass.,  was  attacked  by  some 
fish  that  was  seen  to  overset  and  sink  his  boat  and,  presumably,  devoured 
him,  for  neighboring  fishermen  who  hastened  to  his  rescue  found  no  trace  of 
him.  Whether  his  attacker  was  a  large  shark  or  a  killer  whale  is  an  open 
question. 

"  Murphy  and  Nichols  (Brooklyn  Mus.  Quart.,  vol.  3,  1916,  No.  4,  pp. 
145-160)  give  a  detailed  account  of  this  occurrence. 

»  See  Oudger  (Amer.  Midland  Natural.,  vol.  44,  1050,  p.  714)  for  clinical 
details  of  this  case. 


All  Gulf  of  Maine  sharks  give  birth  to  young 
that  are  not  only  practically  adult  in  structure 
but  of  relatively  large  size  at  birth,  and  there  is  a 
placental  connection  between  mother  and  embryo 
in  some,  but  not  in  others.  Still  other  sharks  lay 
eggs;  this  is  true  of  the  chain  dogfish  {Scyliorhinus 
retifer,  p.  34),  which  is  common  out  on  the  conti- 
nental shelf  from  the  offing  of  Cape  Cod,  south- 
ward, and  of  its  immediate  relatives;  also  of  the 
heterodontids  or  Port  Jackson  sharks  which  are 
not  represented  in  the  Atlantic. 

There  is  so  little  market  for  sharks  in  Gulf  of 
Maine  ports  (attempts  to  introduce  the  dogfish  as 
a  food  fish  having  failed  so  far)  that  the  amounts 
landed  in  Maine  and  Massachusetts  were  only 
about  240,000  pounds  in  1947,  and  about  309,500 
pounds  in  1949;  they  interest  fishermen  chiefly  as 
nuisances  because  of  the  damage  they  do  to  nets 
and  other  gear,  except  that  mackerel  sharks  are 
marketable. 

It  is  possible  to  identify  all  the  sharks  so  far 
known  from  the  Gulf  (and  this  includes  all  that 
are  likely  to  occur  there  except  strays)  bv  the 
sizes  and  relative  locations  of  the  fins,  and  by 
such  tooth  characters  as  may  be  seen  at  a  glance 
at  the  open  mouth  or  easily  felt  with  the  finger 
(after  the  shark  is  dead!). 

We  have  attempted  in  the  following  descriptions 
of  the  several  species  to  include  only  such  features 
as  will  tell  what  shark  is  at  hand ;  for  more  minute 
particulars  we  refer  the  reader  to  our  account  of 
the  sharks  of  the  western  North  Atlantic  (p.  2). 


Key  to  Gulf  of  Maine  Sharks 

1.  There  is  an  anal  fin 2 

There  is  no  anal  fin 16 

2.  Head  greatly  expanded  sidewise,  at  level  of  eyes,  in  hammer-  or  shovel-form 3 

Head  of  ordinary  shape,  with  rounded  or  pointed  snout 4 

3.  Outline  of  front  of  head  only  slightly  concave  opposite  nostrils  if  at  all  so;  grooves  (if  any)  from  nostrils  shorter  than 

horizontal  diameter  of  eyes;  free  tip  of  second  dorsal  fin  is  not  longer  than  forward  margin  of  the  fin;  rear  margin 
of  anal  fin  is  only  weakly  concave;  teeth  near  outer  corners  of  mouth  are  rounded,  without  sharp  cusps. 

Shovel  head,  p.  44 

Outline  of  front  of  head  is  deeply  indented  opposite  each  nostril ;  grooves  from  nostrils  are  more  than  twice  as  long  as 

horizontal  diameter  of  eye;  free  tip  of  second  dorsal  fin  is  considerably  longer  than  front  margin  of  the  fin;  rear 

margin  of  anal  fin  deeply  concave;  teeth  near  corners  of  mouth  are  like  those  near  center  of  mouth,  with  sharp 

cusps Hammerhead,  p.  45 

4.  Caudal  peduncle  (root  of  tail)  is  not  widely  expanded  sidewise  as  a  lateral  keel  on  either  side;  upper  lobe  of  caudal 

fin  is  much  longer  than  lower  lobe 8 

Caudal  peduncle  is  widely  expanded  sidewise  as  a  lateral  keel  on  either  side;  lower  lobe  of  caudal  fin  is  nearly  as  long 
as  upper  lobe,  suggesting  the  caudal  fin  of  a  mackerel  or  swordfish 5 


FISHES   OF  THE  GULF  OF   MAINE  17 

5.  Gill  openings  very  large,  the  first  pair  nearly  meeting  below  the  throat;  teeth  tiny,  many  hundred  in  number;  gill 

arches  with  numerous  horny  gill  rakers  directed  inward-rearward Basking  shark,  p.  28 

Gill  openings,  confined  to  sides  of  head;  teeth  large,  few  in  number;  gill  arches  do  not  have  horny  gill  rakers 6 

6.  Upper  teeth  broadly  triangular,  with  serrate  edges;  anal  fin  is  entirely  behind  second  dorsal  fin 

White  shark,  maneater,  p.  25 

Upper  teeth  with  smooth-edged  cusp,  with  or  without  a  denticle  on  either  side,  at  the  base;  anal  fin  is  not  entirely 

behind  second  dorsal  fin 7 

7.  First  two  teeth  from  center  in  each  jaw  are  similar  to  the  succeeding  teeth;  origin  of  first  dorsal  fin  is  over  or  in  front 

of  inner  corner  of  pectoral  fin  when  latter  is  laid  back ;  forward  part  of  caudal  fin  has  a  small  secondary  lateral  keel 
on  each  side,  below  the  primary  keel  formed  by  the  lateral  expansion  of  the  caudal  peduncle. 

Mackerel  shark,  p.  20 

First  two  teeth  from  center  in  each  jaw  are  noticeably  more  slender  and  more  flexuous  than  the  succeeding  teeth; 

origin  of  first  dorsal  fin  is  behind  inner  corner  of  pectoral  fin  when  latter  is  laid  back;  forward  part  of  caudal  fin  does 

not  have  a  secondary  longitudinal  keel Sharp-nosed  mackerel  shark,  mako,  p.  23 

8.  Upper  lobe  of  caudal  fin  is  nearly  or  quite  as  long  as  head  and  body  combined Thresher,  p.  32 

Upper  lobe  of  caudal  is  less  than  one-half  as  long  as  head  body  combined 9 

9.  Second  dorsal  fin  is  nearly  as  high  vertically  as  first  dorsal  fin 10 

Second  dorsal  fin  is  less  than  one-half  as  high  vertically  as  first  dorsal  fin 12 

10.  First  dorsal  fin  is  wholly  or  mostly  forward  of  the  origin  of  the  pelvic  fins 11 

First  dorsal  fin  is  wholly  posterior  to  bases  of  pelvic  fins Chain  dogfish,  p.  34 

11.  Teeth  high,  narrow,  sharp  pointed,  not  in  mosaic  arrangement;  snout  conical;  fifth  gill  openings  well  in  front  of 

pectoral  fins Sand  shark,  p.  18 

Teeth  small,  low,  rounded,  in  mosaic  arrangement;  snout  flat,  broadly  rounded  in  front;  fifth  gill  openings  are  behind 
origins  of  pectoral  fins Smooth  dogfish,  p.  34 

12.  Origin  of  first  dorsal  fin  far  behind  inner  corner  of  pectoral  fin;  upper  surface  brilliant  blue  in  life. 

Blue  shark,  p.  38 

Origin  of  first  dorsal  fin  is  over  or  anterior  to  inner  corners  of  pectorals;  ground  color  of  upper  surface  is  gray,  brownish 

or  dusky  in  life,  not  bright  blue 13 

13.  Length  of  snout  in  front  of  mouth  is  not  more  than  one-half  as  great  as  breadth  of  mouth;  upper  jaw  has  a  furrow 

on  either  side  extending  from  outer  corner  forward  past  level  of  eye;  caudal  peduncle  with  a  low  longitudinal  keel 
on  either  side;  upper  and  lower  teeth  are  of  shapes  shown  in  figure  11;  their  margins  coarsely  serrate. 

Tiger  shark,  p.  37 

Length  of  snout  in  front  of  mouth  is  more  than  two-thirds  as  gref  t  as  breadth  of  mouth;  furrows  on  upper  jaw,  if 

any,  do  not  extend  forward-inward  as  far  as  level  of  eye;  caudal  peduncle  without  longitudinal  ridges;  teeth  are 

not  of  shape  shown  in  figure  11,  their  margins  either  only  very  finely  serrate  or  smooth 14 

14.  Outer  corners  of  mouth  have  a  short  "labial  furrow"  extending  inward-forward  along  each  jaw;  teeth  are  alike  in 

the  two  jaws,  directed  sharply  outward,  margins  of  upper  teeth  smooth,  as  well  as  those  of  lower  teeth. 

Sharp-nosed  shark,  p.  40 
Outer  corners  of  mouth  have  no  labial  furrow  on  lower  jaw  and  upper  labial  furrow  is  so  short  as  to  be  hardly  notice- 
able; teeth  directed  only  moderately  outward,  their  margins  only  finely  serrate;  lowers  noticeably  more  slender 
than  uppers 15 

15.  Origin  of  first  dorsal  fin  is  about  over  inner  corner  of  pectoral  when  latter  is  laid  back;  vertical  height  of  first 

dorsal  fin  is  less  than  distance  from  eye  to  first  gill  opening Dusky  shark,  p.  41 

Origin  of  first  dorsal  is  about  over  axil  (armpit)  of  pectoral,  its  vertical  height  (after  birth)  is  at  least  as  great  as 
distance  from  eye  to  third  gill  opening Brown  shark,  p.  43 

16.  Trunk  much  flattened  dorso-ventrally;  eyes  on  top  of  head;  front  margins  of  pectorals  overlap  the  gill  openings. 

Angel  shark,  note,  p.  18 
Trunk  subcylindrical;  eyes  on  side  of  head;  front  margins  of  pectorals  do  not  overlap  the  gill  openings    17 

17.  Each  dorsal  fin  is  preceded  by  a  stout  and  conspicuous  spine 18 

Doisal  fin-spines  either  lacking,  or  are  so  nearly  concealed  in  the  skin  that  their  presence  can  be  detected  by  touch 

only 20 

18.  Upper  teeth  with  5  erect  cusps;  lower  teeth  with  only  one  cusp,  the  successive  cusps  directed  outward,  forming  a 

nearly  continuous  horizontal  cutting  edge  all  along  the  jaw Etmopterus  princeps,  p.  47 

Upper  and  lower  teeth  are  alike  in  shape 10 

19.  Upper  teeth  quadrangular  as  well  as  lower  teeth,  with  one  cusp  directed  outward,  forming  a  nearly  continuous 

horizontal  cutting  edge  along  each  jaw Spiny  dogfish,  p.  47 

Upper  and  also  lower  teeth  each  have  3  to  5  erect,  triangular  cusps Black  dogfish,  p.  51 

20.  First  dorsal  fin  well  in  advance  of  pelvic  fins;  upper  teeth  noticeably  different  in  shape  from  lower  teeth 21 

First  dorsal  fin  stands  over  posterior  part  of  bases  of  pelvic  fins;  upper  are  teeth  similar  to  lower  teeth  in  shape. 

Bramble  shark,  d.  56 


18 


FISHERY  BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


21.  Lower  teeth  erect,  triangular,  their  edges  serrate Dalatias  licha,  p.  55 

Lower  teeth  quadrate,  the  cusp  directed  outward,  forming  a  nearly  continuous  horizontal  cutting  edge;  their  outer 

margins  deeply  notched,  the  edges  smooth . 22 

22.  Dermal  denticles  rounded,  overlapping,  scale-like,  entirely  concealing  the  skin  (fig.  20) ;  each  dorsal  fin  is  preceded 

by  a  short  spine,  embedded  nearly  to  its  tip  in  the  skin,  but  recognizable  by  touch Portuguese  shark,  p.  52 

Dermal  denticles  conical,  only  moderately  close  set,  the  skin  visible  between  them;  dorsal  fins  not  preceded  by 

spines Greenland  shark,  p.  53 

Note. — Not  yet  known  from  the  Gulf  of  Maine  though  reported  from  Marthas  Vineyard. 

THE  SAND  SHARKS.     FAMILY  CARCHARIIDAE 


Outstanding  characteristics  of  the  sand  sharks 
are  that  they  have  an  anal  fin ;  the  two  dorsal  fins 
are  without  spines  and  are  nearly  equal  in  size ;  the 
rear  end  of  the  base  of  the  first  dorsal  is  over  or  in 
front  of  the  origin  of  the  pelvic  fins;  the  anal  fin  is 
about  as  large  as  the  dorsals;  the  upper  lobe  of  the 
caudal  fin  is  much  longer  than  the  lower,  but 
occupies  not  more  than  one-third  of  the  total  length 
of  the  fish;  there  are  no  lateral  keels  on  the  caudal 
peduncle;  the  fifth  gill  openings  are  farther  forward 
than  the  origins  of  the  pectoral  fins;  and  the  teeth 
are  slender  and  sharp-pointed. 

Sand   shark  Carcharias  taurus  Rafinesque  1810 

Dogfish  shark;  Ground  shark 

Bigelow  and  Schroeder,  1948,  p.  100. 
Garman,  1913,  pi.  6,  figs.  1-3. 

Description.  — The  large  size  of  the  second  dorsal 
fin,  and  of  the  anal  as  well  (which  is  about  equal  to 
the  first  dorsal  instead  of  much  smaller)  is  of  itself 
enough  to  distinguish  this  species  from  all  other 


Gulf  of  Maine  sharks.  The  fact  that  the  first  dorsal 
fin  is  located  but  little  in  front  of  the  pelvics,  and 
that  the  trunk  seems  crowded  with  fins  of  equal 
size,  is  a  useful  field  mark.  We  may  also  point  out 
that  the  pectoral  fins  are  not  much  larger  than  the 
other  fins — triangular  rather  than  sickle-shaped; 
that  the  upper  lobe  of  the  tail  is  nearly  one-third 
as  long  as  head  and  body  together  and  notched 
near  its  tip,  with  the  lower  lobe  about  one-third 
as  long  as  the  upper  lobe;  and  that  the  head  is 
flat  above,  the  snout  short,  conical  with  rather 
sharp  tip.  The  teeth  also  (alike  in  the  two  jaws) 
are  diagnostic,  being  long,  narrow,  sharp-pointed, 
and  smooth-edged,  with  one  (rarely  two)  small 
spurs  ("denticles")  on  either  side  near  the  base. 

Size. — Most  of  the  sand  sharks  that  are  caught 
in  the  northern  part  of  their  American  range,  from 
Delaware  Bay  to  Cape  Cod,  are  immature,  of 
perhaps  4  to  6  feet.  But  adults  up  to  8  or  9  feet 
long  are  reported  there  from  time  to  time,  espe- 
cially from  the  vicinity  of  Nantucket,  where  a 
commercial  shark  fishery  yielded  many  of  them  in 


Figure  4. — Sand  shark  (Carcharias  taurus),  about  40  inches  long,  Cape  Cod;  and  upper  and  lower  teeth  from  front  part 
of  mouth  of  a  larger  specimen  from  New  Jersey,  about  natural  size.  From  Bigelow  and  Schroeder.  Drawings  by 
E.  N.  Fischer. 


FISHES   OF  THE  GULF  OF   MAINE 


19 


the  early  1920's.  And  large  ones,  alone,  have 
been  reported  from  North  Carolina,  southward. 
The  greatest  recorded  length  is  10  feet  5  inches, 
from  southwestern  Florida.  And  the  sand  shark 
does  not  mature  sexually  until  perhaps  7  feet  long, 
or  more.  A  weight  of  250  pounds  is  recorded  for 
one  8  feet  10  inches  long,  showing  how  much 
lighter  a  fish  this  is,  length  for  length,  than  various 
other  sharks. 

Color.- — Light  gray-brown  above,  darkest  along 
back,  snout,  and  upper  sides  of  pectorals,  paling  on 
the  sides  to  grayish  white  on  lower  surface;  sides 
of  trunk  rearward  from  pectorals  variously  marked 
with  roundish  to  oval  spots,  of  which  there  may  be 
upwards  of  100,  varying  in  color  from  yellowish 
brown  to  ocher  yellow.  The  rear  margins  of  the 
fins  are  edged  with  black  on  some  specimens, 
but  not  on  others. 

Habits  and  food. — Despite  its  trim  appearance 
and  voracious  appetite,  this  is  a  comparatively 
sluggish  shark,  living  mostly  on  bottom  or  close  to 
it;  more  active  and  taking  a  bait  more  freely  at 
night  than  by  day.  During  its  summer  visits  to 
the  New  England  coast  it  holds  so  close  to  the 
coast  that  it  has  never  been  reported  from  Georges 
Bank,  or  from  the  outer  part  of  the  Continental 
Shelf.  Most  of  those  caught  are  from  depths  not 
greater  than  1  to  5  fathoms,  occasionally  perhaps 
as  deep  as  10  fathoms,  and  many  come  right  in  to 
tide  line  along  the  beaches.  They  may  sometimes 
be  seen  moving  slowly  to  and  fro  at  the  surface, 
over  bars,  with  dorsal  and  caudal  fins  showing 
above  the  water;  and  they  sometimes  enter  the 
mouths  of  rivers.  They  capture  great  numbers  of 
small  fish,  which  are  their  chief  diet,  particularly 
menhaden,  cunners,  mackerel,  skates,  silver  hake, 
flounders,  alewives,  butterfish,  and  south  of  Cape 
Cod,  scup,  weakfish,  and  bonito.  Sand  sharks 
have  been  seen  surrounding  and  harrying  schools  of 
bluefish ;  they  have  even  been  known  to  attack  nets 
full  of  bluefish,  which  gives  a  measure  of  their 
voracity.     They  also  eat  lobsters,  crabs,  and  squid 

Breeding. — -The  eggs  of  the  sand  shark  are 
hatched  within  the  parent  and  are  retained  there 
until  the  resultant  young  are  ready  for  independent 
existence,  but  there  is  no  placental  connection 
between  mother  and  developing  embryo.  It  has 
recently  been  discovered  that  while  a  ripe  female 
contains  a  large  number  of  eggs,  only  two  embryos 
develop  as  a  rule,  one  in  each  oviduct;  they  are 
nourished   (at  least   largely)   by  swallowing   the 


unfertilized  eggs  M  with  which  the  stomach  of  the 
embyro  becomes  greatly  distended.  Females 
with  large  embryos  have  so  far  been  reported  only 
from  Florida  and  from  Louisiana,  whereas  others 
taken  near  Woods  Hole  have  contained  eggs  only, 
making  it  likely  that  the  small  specimens  that  are 
so  common  along  southern  New  England  have 
come  from  a  more  southerly  birthplace. 

General  range. — Coastal  waters  on  both  sides  of 
the  Atlantic;  Maine  to  Florida  and  Brazil  in  the 
west;  Mediterranean,  tropical  West  Africa,  Ca- 
naries, and  Cape  Verdes  in  the  east;  also  South 
Africa ;  represented  in  Argentine  waters  and  in  the 
Indo-Pacific  by  close  relatives. 

Occurrence  in  the  Gulf  of  Maine. — The  sand 
shark  is  by  far  the  most  common  of  its  tribe,  next 
to  the  smooth  and  spiny  dogfishes,  along  southern 
New  England  and  at  the  westerly  entrance  to  the 
Gulf  of  Maine.  It  is  plentiful  at  Woods  Hole  from 
June  to  November,  to  be  found  anywhere  in  that 
region  in  shoal  waters,  even  coming  up  to  the 
wharves.  At  Nantucket,  too,  it  is  so  abundant 
that  shark  fishing,  with  the  sand  shark  as  the  chief 
objective,  is  a  popular  sport.  The  facts  that  a 
catch  of  about  1,900  sharks  by  three  boats  on 
Horseshoe  Shoal,  in  Nantucket  Sound,  June  to 
September  1918,  was  mostly  of  this  species,  as 
was  another  catch  of  350  sharks,  taken  near  Nan- 
tucket in  the  early  1920's,  illustrate  their  numbers 
there.  Scattered  sand  sharks  are  also  caught  along 
the  outer  beaches  of  Cape  Cod  by  surf  anglers 
(published  records  are  for  Monomoy,  Chatham, 
and  Provincetown)  and  there  are  enough  of  them 
along  this  stretch  of  beach  in  some  summers  (1951 
was  a  case  in  point)  for  them  to  be  a  nuisance  to 
anglers  casting  for  striped  bass  in  the  surf  at  night. 

In  August  1947  we  saw  a  large  one  at  the  surface 
pursuing  a  striped  bass,  that  was  being  hauled 
aboard  a  fishing  boat  on  a  hand  line,  in  the 
eastern  side  of  Cape  Cod  Bay,  where  fishermen 
tell  us  that  this  is  not  an  unusual  happening.  But 
this  appears  to  be  the  northern  boundary  to  their 
occurrence  in  any  numbers,  or  with  regularity. 
True,  they  are  recorded  at  Cohasset,  on  the  south- 
ern shore  of  Massachusetts  Bay,  where  we  caught 
one  about  4  feet  long,  years  ago  in  Boston  Bay, 
and  at  Lynn,  Mass.  But  so  rarely  does  it  stray 
north  of  Cape  Ann  that  it  has  been  reported  only 


»  For  an  account  of  the  embryos,  see  Springer,  Copeia,  1948,  No.  3,  pp. 
153-156. 


20 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


twice  from  Casco  Bay,  and  once  from  St.  Andrews, 
New  Brunswick,  near  the  mouth  of  the  Bay  of 
Fundy,  its  most  northerly  known  outpost,  where 
one  was  taken  in  a  weir  in  1913. 

In  New  England  waters  the  sand  shark  occurs 
only  as  a  summer  visitor.  The  winter  home  of 
those  that  summer  along  the  northeastern  United 
States  is  not  known,  nor  has  any  increase  been 
noted  in  Florida  waters  (where  they  are  taken  at 
all  times  of  year)  coincident  with  their  winter  dis- 
appearance from  the  northern  part  of  their  range. 
Like  various  bony  fishes  they  may  move  offshore, 
and  perhaps  southward,  to  escape  winter  chilling. 

Importance. — There  were  commercial  fisheries 
for  the  sand  shark  around  Nantucket  during  the 
first  quarter  of  the  present  century,  but  these  were 
short  lived,  reputedly  because  of  exhaustion  of  the 


local  stock.  And  the  sand  shark  is  of  no  commer- 
cial importance  on  the  New  England  coast  at 
present.  Westward  from  Cape  Cod  it  is  of  some 
interest  to  anglers,  who  catch  considerable  num- 
bers, both  as  objects  of  special  pursuit,  for  it  takes 
almost  any  natural  bait  readily,  or  incidentally 
while  surf  casting  for  better  fish.  But  it  is  not 
plentiful  enough  in  the  Gulf  of  Maine  to  be  worth 
fishing  for. 

There  is  no  record  of  attacks  by  sand  sharks  on 
human  beings  in  North  American  waters,  though 
bathers  often  come  close  to  them.  Our  own  experi- 
ence bears  this  out;  in  fact,  it  is  looked  upon  as  a 
harmless  nuisance  on  the  New  England  coast 
wherever  it  is  plentiful  enough  to  be  familiar. 
But  its  relative  (or  relatives)  of  East  Indian  waters 
have  a  more  sinister  reputation. 


MACKEREL  SHARKS.     FAMILY  ISURIDAE 


Sharks  of  this  family  are  easily  recognizable 
by  the  very  firm  half-moon-shaped  (technically 
lunate)  caudal  fin,  with  lower  lobe  but  little  shorter 
than  the  upper,  in  combination  with  large  awl-like 
or  blade-shaped  teeth,  and  with  gill  openings 
larger  than  any  other  Gulf  of  Maine  shark  except 
the  basking  shark.  Their  tail  fins,  in  fact,  recall 
the  tails  of  such  bony  fishes  as  the  mackerel  tribe 
or  the  swordfish,  in  outline,  likewise  in  firm  tex- 
ture, hence  their  common  name.  The  basking 
shark  also  has  a  caudal  fin  and  peduncle  of  this 
same  sort,  but  its  teeth  are  minute  and  very 
numerous,  and  its  gill  openings  are  so  long  that 
those  of  the  two  sides  nearly  meet  on  the  lower 
surface  of  the  throat. 

Other  diagnostic  features  are  that  they  have  an 
anal  fin ;  that  their  caudal  peduncle  is  expanded  as 
a  prominent  longitudinal  keel  on  either  side;  that 
their  dorsal  fins  are  not  preceded  by  spines;  and 
that  the  inner  margins  of  their  gill  arches  do  not 
have  horny  gill  rakers. 

Mackerel  shark  Lamna  nasus  (Bonnaterre)  1788 

Porbeagle;    Blue    dog    (in    Gulf  of  Maine) 

Bigelow  and  Schroeder,  1948,  p.  112. 

Garman,  1911,  pi.  6,  figs.  4-6  (as  Isurus  ■punctatus). 

This  is  a  stout,  heavy-shouldered  shark,  tapering 
in  front  to  a  pointed  conical  snout  and  behind  to 
a  very  slim  tail  root.  Its  dorsal  and  pectoral  fins 
are  large;  the  former,  originating  a  little  rearward 


of  the  armpits  of  the  pectorals,  is  triangular  and 
about  as  high  as  it  is  long;  the  pectoral  fins  are 
only  half  as  broad  as  long.  The  second  dorsal  and 
anal  fins  are  very  small  indeed,  and  the  pelvics 
but  little  larger.  The  second  dorsal  fin  stands 
over  the  anal.  There  is  a  conspicuous  transverse 
furrow  or  pit  on  the  upper  surface  of  the  root  of  the 
tail,  also  one  on  the  lower  surface  close  in  front  of 
the  origin  of  the  caudal  fin.  The  lower  lobe  of  the 
caudal  fin  is  two-thirds  to  three-fourths  as  long 
as  the  upper  lobe,  and  there  is  a  small  secondary 
keel  on  the  base  of  the  caudal  fin  on  either  side, 
below  and  behind  the  rear  end  of  the  primary 
keel  formed  by  the  sidewise  expansion  of  the 
caudal  peduncle. 

The  teeth  of  the  porbeagle  are  alike  in  the  two 
jaws,  slender,  pointed,  smooth-edged,  and  with  a 
sharp  denticle  near  the  base  on  each  side  (young 
fish  may  not  have  these)  which  the  mako  lacks 
(P-  23). 

The  only  Gulf  of  Maine  sharks  with  which  the 
porbeagle  might  be  confused  are  the  maneater 
(p.  25),  or  the  mako  (p.  23).  And  it  is  easily 
told  from  the  former  by  its  slender,  smooth-edged 
teeth,  as  well  as  by  the  position  of  its  second 
dorsal  fin  directly  over  the  anal;  from  the  mako 
by  the  shape  of  its  teeth  (cf.  fig.  5  with  fig.  6), 
each  usually  with  a  small  basal  denticle  on  either 
side,  which  the  mako  lacks;  also  by  its  stouter 
body  and  by  the  presence  of  the  secondary 
longitudinal  keel  on  the  anterior  part  of  its 
caudal  fin. 


FISHES    OF   THE   GULF   OF    MAINE 


21 


Figure  5. — Mackerel  shark  (Lamna  nasus),  about  37  inches  long,  Nahant,  Massachusetts.  Upper  and  lower  first  to 
fifth  teeth  from  center  of  jaw  of  a  larger  specimen  from  Platts  Bank,  about  0.7  times  natural  size.  From  Bigelow 
and  Schroeder.     Drawings  by  E.  N.  Fischer. 


Color. — Dark  bluish  gray  to  bluish  black  above, 
including  the  upper  surfaces  of  the  pectorals, 
changing  abruptly,  low  down  on  the  sides,  to  white 
below;  lower  surfaces  of  pectorals  dusky  to  black 
on  the  outer  one-half  to  one-third,  more  or  less 
mottled  white  and  dark  toward  their  bases,  and 
with  the  anterior  and  posterior  edges  narrowly 
rimmed  with  black;  the  anal  is  white  or  slightly 
dusky. 

Size. — The  common  run  of  mackerel  sharks  in 
the  Gulf  of  Maine  are  from  4  to  6  feet  long,  with 
few  heavier  than  200  pounds;  thus  18  recently 
landed  at  Portland  and  Eastport,  Maine,31  averaged 
4  feet  5  inches,  the  largest  being  about  8  feet  long, 
the  smallest  3  feet  7  inches. 

Specimens  longer  than  7  to  8  feet  are  not 
common;  only  two  longer  than  8  feet  have  been 
recorded  previously  from  the  Gulf  of  Maine,  one 
of  which  was  10  feet,32  the  largest  recorded 
from  either  side  of  the  North  Atlantic.  This  shark 
has  been  said  to  reach  a  length  of  12  feet.  But 
the  sizes  of  sharks  often  are  overstated,  unless 
actually  measured,  point  to  point,  not  around  the 
curve  of  the  body.  Information  as  to  the  relation- 
ship between  length  and  weight  is  restricted  to  a 
report  of  305  pounds  at  8  feet  3  inches,  and  of 
about  400  pounds  at  about  9  feet.  One  3  feet  long 
that  we  measured  weighed  20  pounds. 

»  8cattergood,  Copela,  1949,  No.  1,  pp.  71-72. 
n  Hubbs,  Copela,  No.  173,  19?3,  p.  101 


Habits. — The  whole  mackerel-shark  tribe  lead  a 
pelagic  life,  wandering  about  over  the  ocean  in 
pursuit  of  the  fishes  on  which  they  prey,  and 
often  uniting  in  small  companies,  though  they 
can  hardly  be  called  gregarious.  Like  swordfish 
they  spend  much  time  at  the  surface  on  calm  days, 
when  their  triangular  back  fins,  followed  by  the 
tip  of  the  caudal  fin  (the  bluntness  of  the  former 
and  the  wavy  track  of  the  latter  identify  the 
shark  as  such)  may  often  be  seen  cutting  through 
the  water.  We  have  sailed  close  to  sharks  probably 
of  this  species  again  and  again,  only  to  see  them 
sound,  just  out  of  harpoon  range,  plainly  visible 
at  first  but  soon  fading  from  sight  as  they  swim 
downward. 

The  porbeagle  has  often  been  described  as 
active  and  strong  swimming.  But  it  puts  up 
only  a  very  feeble  resistance  when  hooked. 
We  have  never  seen  or  heard  of  one  jumping,  as 
the  mako  often  does  (p.  24),  nor  is  there  any 
difficulty  in  landing  one  of  4  to  5  feet  on  an 
ordinary  cod  line.  It  is,  in  fact,  as  proverbial 
among  fishermen  for  its  sluggishness  when  hooked, 
as  is  the  mako  for  its  activity.  While  often  seen 
"finning,"  many  are  caught  close  to  the  bottom, 
in  depths  down  to  80  fathoms  in  the  gill  net 
fishery  for  ground  fish  that  is  carried  on  from 
Portland,  Maine;  some  also  on  bottom  on  cod 
lines;  how  much  deeper  they  may  descend  is 
not  known. 


22 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Food. — la  the  Gulf  of  Maine  the  porbeagle  feeds 
chiefly  on  mackerel  and  on  the  herring  tribe;  on 
butterfish;  on  ground  fish,  as  cod,  hake,  cusk, 
rosefish,  flounders,  or  other  kinds  available;  and 
on  squid.  It  has  also  the  annoying  custom  of 
foraging  on  the  cod  and  other  fish  that  have  been 
hooked  on  long  lines  and  biting  off  the  snoods. 
It  is  also  known  to  prey  on  the  spiny  dogfish  in 
the  eastern  Atlantic;  probably  in  the  Gulf  of 
Maine  also.  But  we  find  no  record  of  its  eating 
crustaceans  of  any  kind. 

Breeding. — The  mackerel  shark  tribe  are  ovovi- 
viparous;  that  is,  the  eggs  are  hatched  within  the 
maternal  oviducts,  but  there  is  no  placental  con- 
nection between  mother  and  young.  The  embryos, 
like  those  of  the  sand  shark  (p.  19),  are  nourished 
chiefly  by  swallowing  the  unfertilized  eggs  that 
lie  nearby  in  the  "uterus,"  and  their  stomachs 
become  enormously  swollen  by  the  masses  of  yolk 
that  are  eaten  in  this  way.  Another  interesting 
feature  of  the  porbeagle  embryo  is  that  the  upper 
lobe  of  its  caudal  fin  is  much  longer  at  first  than 
the  lower  lobe,  the  latter  increasing  in  relative 
length  with  growth.  The  embryos  also  are  very 
large  at  birth;  young  of  18,  19,  and  24  inches  have, 
for  example,  been  found  in  a  five-foot  mother. 
Corresponding  to  their  large  size,  gravid  females 
contain  only  one  to  four  young  (0-2  in  each 
oviduct) . 

General  range. — Continental  waters  in  both  sides 
of  the  North  Atlantic;  southern  Scandinavia, 
Orkneys  and  North  Sea  southward  to  the  Mediter- 
ranean and  northwest  Africa  in  the  east;  northern 
coast  of  Newfoundland,33  Newfoundland  Banks 
and  Gulf  of  St.  Lawrence  to  New  Jersey  and  per- 
haps to  South  Carolina  in  the  west;  represented 
in  the  northwest  Pacific  and  in  Australian-New 
Zealand  waters  by  forms  that  are  closely  allied  to 
it,  but  not  identical. 

Occurrence  in  the  Gulf  of  Maine. — It  has  been 
known  from  the  days  of  the  earliest  settlement 
that  stout-shouldered,  surface-swirnming  sharks 
of  moderate  size,  with  "mackerel"  tails  and  slen- 
der, smooth-edged  teeth  are  tolerably  common  in 
the  Gulf  of  Maine;  they  are  universally  referred  to 
by  the  fishing  population  as  "mackerel  sharks." 
During  the  first  half  of  the  last  century  only  one 
such  shark  species  was  recognized  in  our  waters. 
And  while  more  recent  researches  have  proved 

••  One  reported  at  Raleigh,  on  the  Newfoundland  side  of  the  Strait  of 
Belle  Isle,  July  1929,  by  Dr.  W.  O.  Jeflers. 


that  two  actually  occur  within  the  limits  of  the 
Gulf  (this  and  the  next  described)  the  present 
species  is  the  more  northerly  of  the  pair,  and 
much  the  more  frequently  taken  in  the  Gulf. 
Hence  it  is  probable  that  most  of  the  mackerel 
sharks  that  fishermen  often  see  swimming  lazily 
on  the  surface,  and  often  catch,  off  the  shores  of 
northern  New  England,  belong  here. 

Seemingly,  the  chief  centers  of  population  for 
the  porbeagle  in  the  western  Atlantic  are  along 
outer  Nova  Scotia,  and  in  the  western  side  of  the 
Gulf  of  Maine.  Thus,  while  there  are  but  two 
published  records  for  it  from  the  Newfoundland 
Banks,  and  one  (besides  verbal  reports)  in  the 
Gulf  of  St.  Lawrence,  fishermen  report  it  as  the 
commonest  large  shark  along  the  Atlantic  coast 
of  Nova  Scotia.  Apparently  it  tends  to  shun  the 
cold  waters  of  the  Bay  of  Fundy,  for  it  is  recorded 
only  twice  from  Passamaquoddy  Bay,  one  in 
August  1900,  the  other  on  October  3,  1935.34  But 
it  is  so  plentiful  farther  west  in  the  Gulf  that  inci- 
dental catches  are  on  record  of  19  that  were  taken 
in  one  night  by  six  men  on  hand  lines,  and  of  about 
150  taken  by  one  crew  during  three  weeks'  cod 
fishing  near  Monhegan  Island,  Maine.  We  have 
ourselves  hooked  or  sighted  about  one  per  three 
or  four  days'  fishing,  on  the  cod  grounds  in  general 
in  the  western  side  of  the  Gulf,  the  majority  near 
Platts  Bank  off  Cape  Elizabeth,  but  some  also  on 
Nantucket  Shoals.36  Certainly  it  is  the  most  often 
seen  of  the  larger  sharks  around  the  Isles  of  Shoals 
and  near  Cape  Ann,  and  it  has  been  characterized 
repeatedly  as  "common"  in  Massachusetts  Bay.38 

To  the  westward  the  porbeagle  is  described  as 
not  uncommon  near  Woods  Hole  (we  have  not 
seen  it  there).  We  saw  a  small  one  about  3  feet 
long  taken  in  an  otter  trawl  at  60  fathoms,  off 
Marthas  Vineyard,  on  February  20,  1950,  by  the 
Eugene  H;  and  it  has  been  reported  on  several 
occasions  from  Rhode  Island  waters.  But  it  ap- 
pears only  as  a  stray  off  New  York  and  to  the 
southward. 

Thus,  the  latitudinal  range  within  which  it 
occurs  regularly  off  the  American  coast  covers 
only  something  like  5°.  And  its  on-  and  offshore 
range  is  correspondingly  so  narrow  that  no  report 

'<  Reported  by  McOonigle  and  Smith,  Proc.  Nova  Scotia  Inst.  Sci.,  vol. 
19, 193S,  p.  160. 

11  Cod  tagging  cruises  of  the  U.  S.  Bureau  of  Fisheries. 

18  Actually  no  sharks  other  than  the  spiny  dogfish  (p.  47)  are  "common" 
In  the  Gulf  of  Maine,  In  the  sense  that  this  term  Is  applied  to  such  fish  as 
herring,  cod,  mackerel,  and  other  species,  but  only  as  relative  to  other  sharks 
of  corresponding  sizes. 


FISHES   OF  THE  GULF  OF   MAINE 


23 


of  it  has  come  to  hand  from  Georges  or  Browns 
Banks,  only  one  from  the  Nova  Scotia  slope  off 
Sable  Island,  and  two  from  the  Grand  Banks,  as 
just  noted.  On  the  other  hand,  few  come  in-shore 
close  enough  to  be  picked  up  in  pound  nets  or 
weirs. 

All  published  records  of  mackerel  sharks  from 
the  Gulf,  and  all  that  we  have  seen  there,  have 
been  in  the  warm  half  of  the  year,  and  something 
like  70  percent  of  the  landings  of  porbeagles  on 
the  coast  of  Maine  are  for  August  to  November. 
But  its  presence  in  the  Gulf  in  winter  is  proved 
by  our  receipt  of  a  photograph  of  a  porbeagle 
embryo,  taken  from  a  female  caught  in  January, 
off  Portland,  Maine,  in  1927.  And  it  is  also 
caught  in  winter  as  well  as  in  summer  in  north 
European  waters.  Apparently  it  simply  descends 
into  deeper  water  during  the  winter  to  escape  low 
surface  temperatures,  feeding  little,  else  more  of 
them  would  have  been  caught  in  the  Gulf  during 
the  winter  fishery  with  long  lines  for  hake  (llro- 
phycis) . 

In  the  Gulf  of  Maine,  females  containing  em- 
bryos have  been  taken  in  August  (near  Monhegan 
Island,  Maine) ;  in  October  (off  Barnstable,  Mass.) ; 
in  November  (off  Portland,  Maine) ;  and  in  Jan- 
uary (off  Portland,  Maine).  But  the  fact  that  the 
largest  embryos  have  been  found  in  European  seas 
in  summer  suggests  that  most  of  the  young  are 
not  born  until  then. 

Importance. — The  liver  oil  of  the  porbeagle, 
mixed  with  other  fish  oils,  was  in  demand  for  use 
in  tanning  leather  during  the  first  quarter  of  the 
19th  century.  And  it  is  interesting  to  read  that 
as  much  as  1 1  gallons  of  oil  has  been  obtained  from 
the  liver  of  a  single  shark  9  feet  long. 

This  demand  had  almost  entirely  died  before 
1850  and  has  never  revived.  But  a  new  demand 
has  developed  of  late  years  for  porbeagle  meat, 
which  resembles  swordfish  in  taste  as  well  as  in 
appearance,  resulting  in  landings  for  this  purpose 
of  about  46,000  pounds  in  1944  on  the  coast  of 
Maine,  and  of  71,600  pounds  in  1945.  Assuming 
an  average  weight  of,  say,  50  pounds,  this  corre- 
sponds to  a  commercial  catch  of  about  900  to  1,400 
sharks.  There  is  no  special  fishery  for  porbeagles 
at  present  in  the  Gulf  of  Maine,  or  for  any  other 
sharks  for  that  matter.  About  four-fifths  of  those 
brought  in  are  taken  in  gill  nets  set  on  bottom  for 
ground  fish,  and  most  of  the  sharks  caught  in  this 
way  are  landed  in  Portland,  Maine.     The  re- 


mainder are  taken  by  seines,  traps,  weirs,  hook  and 
line  or  harpoons.  And  most  of  the  porbeagles 
taken  in  these  ways  are  discarded  at  sea.37  The 
porbeagle  is  not  "game"  enough  to  be  of  any  in- 
terest to  sport-anglers. 

Sharp-nosed  mackerel  shark  Isurus  oxyrinchus 
Rafinesque  1810 

Atlantic  mako 
Bigelow  and  Schroeder,  1948,  p.  124. 

Description. — This  shark  resembles  the  common 
mackerel  shark  so  closely  that  we  need  merely 
point  out  the  points  of  difference.  Most  obvious 
of  these  is  that  while  the  first  dorsal  originates 
about  above  the  armpits  of  the  pectorals  in  the 
common  mackerel  shark,  it  stands  over  or  behind 
the  inner  corner  of  the  pectoral  in  the  mako,  and 
that  the  second  dorsal  originates  a  short  distance 
in  front  of  the  anal.  The  teeth,  too,  differ  rather 
noticeably  in  appearance,  for  while  of  the  same 
awl-like  type,  those  of  the  mako  lack  the  lateral 
spurs  or  denticles  that  are  characteristic  of  all  but 
the  smallest  porbeagles,  and  those  in  the  front  part 
of  the  mouth  are  conspicuously  flexuous  in  form. 
The  mako,  too,  is  more  slender  bodied;  its  snout  is 
more  narrowly  conical ;  its  upper  and  lower  caudal 
lobes  are  more  nearly  equal  in  length;  and  the 
forward  part  of  its  caudal  fin  lacks  the  secondary 
lateral  keels  that  are  to  be  seen  on  the  caudal  fin 
of  the  porbeagle  (cf.  fig.  6  with  fig.  5). 

Color. — Deep  blue-gray  above  when  fresh- 
caught,  appearing  cobalt  or  ultramarine  in  the 
water,  with  gradual  transition  along  the  sides  to 
snow-white  below;  but  turning  dark  slate  gray 
above  soon  after  death  (especially  if  preserved), 
and  to  bluish  white  or  pale  dirty  gray  below  and 
on  the  lower  surfaces  of  the  pectorals. 

Size. — The  maximum  length  reported  for  a  spec- 
imen of  the  Atlantic  mako  that  was  actually  meas- 
ured is  about  12  feet,38  though  it  has  been  said  to 
grow  to  13  feet.  The  largest  western  Atlantic 
specimen  of  which  we  find  definite  record,  taken 
off  St.  Petersburg,  Fla.,  was  10  feet  6  inches  long, 
and  one  nearly  as  large  (10  ft.  2  in.)  was  caught  off 
New  York  Harbor  many  years  ago.  But  the  com- 
mon run  caught  off  the  middle  Atlantic  United 

•'See  Scattergood,  Copela,  1949,  p.  70,  for  further  details  as  to  landings  In 
Maine  and  methods  of  capture. 
■  3.7  meters  as  calculated  from  the  size  of  Its  jaws. 


24 


FISHERY   BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


Figube  6. — Sharp-nosed  mackerel  shark,  or  Mako  (Isurus  oxyrinchus),  about  64J^  inches  long,  Maryland.     Below,  teeth 
in  front  of  mouth  of  a  large  specimen,  Cape  Cod.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


States  are  perhaps  5  to  8  feet  long.  Males  of 
about  6  feet  are  sexually  mature  (as  indicated  by 
the  claspers).  Recorded  weights  at  different 
lengths  are  about  135  pounds  at  6  feet,  230  pounds 
at  7  feet  8  inches;  and  about  300  pounds  at  8  feet. 
The  heaviest  Atlantic  mako  caught  on  rod  and  reel 
of  which  we  have  found  record  was  one  of  786 
pounds  taken  off  Bimini,  Bahamas,  by  Ernest 
Hemingway  in  1936;  the  largest  Pacific  mako  one 
of  798  pounds,  taken  by  E.  White-Wickham  off 
New  Zealand.39 

Habits. — This  is  one  of  the  most  active  and 
swift  swimming  of  the  sharks.  In  seas  where  it 
is  more  common  than  it  is  in  our  Gulf,  it  is  often 
seen  swimming  at  the  surface,  and  it  is  famous 
for  its  habit  of  leaping  clear  of  the  water,  not 
only  when  hooked,  but  under  natural  conditions. 
Seemingly  it  preys  chiefly  on  schools  of  smaller 
fishes  of  the  mackerel  and  herring  tribes.  But  it 
also  attacks  larger  fishes.  A  730-pound  mako,  for 
example,  that  was  harpooned  near  Bimini  in  the 
Bahamas,  contained  a  120-pound  swordfish 
(Xiphias  gladius)  almost  entire,  while  one  weighing 
about  800  pounds,  harpooned  off  Montauk,  Long 
Island,  was  seen  attacking  a  swordfish,  and  was 

"  A  Soutb  African  shark  of  2,176  pounds,  landed  on  rod  and  reel,  and  re- 
ported as  a  mako.  Is  proved  by  the  photograph  of  Its  teeth  (London  Illus. 
News,  July  14, 1628,  p.  83)  to  hare  been  a  maneater  (Corcharodon). 


found  when  landed  to  contain  a  large  amount  of 
its  flesh* 

Young  embryos  of  the  mako,  like  those  of  the 
porbeagle  (p.  22),  have  greatly  dilated  stomachs, 
being  nourished  on  the  unfertilized  eggs  that  he 
near  them  in  the  oviducts,  and  they  are  very 
large  at  birth,  relative  to  the  size  of  the  mother. 

General  range. — This  is  an  oceanic  shark,  of  the 
tropical  and  warm-temperate  belts  of  the  Atlantic 
north  and  south,  including  the  Mediterranean  in 
the  east  and  the  Caribbean  and  Gulf  of  Mexico 
in  the  west.  It  is  represented  in  the  corresponding 
thermal  belts  of  the  Pacific  and  Indian  Oceans  by 
a  close  ally,  the  Pacific  mako  Isurus  glaucus. 

Occurrence  in  the  Gulf  oj  Maine. — The  center  of 
abundance  for  the  mako  lies  in  warmer  seas  to 
the  southward  of  our  Gulf.  Considerable  num- 
bers journey  northward,  however,  in  summer 
along  the  continental  shelf,  as  far  as  to  the  offing 
of  southern  New  England,  and  a  few  are  caught 
off  Woods  Hole.  One  of  the  earliest  accounts  of  it 
in  American  waters  was  based  partly  on  one  from 
Cape  Cod.  During  the  past  few  summers  we 
have  heard  repeatedly  of  makos  seen  jumping,  or 
occasionally   hooked   near   the  northern   end   of 

•  See  Fairlngton  (Field  and  Stream,  vol.  47,  Feb.  1943)  for  these  Instances 
of  the  mako  attacking  swordfish,  and  for  other  Interesting  notes  on  this 
shark. 


FISHES   OF   THE   GULF   OF    MAINE 


25 


Cape  Cod,  and  in  the  summer  of  1941  one  about 
six  feet  long  was  landed  on  rod  and  reel  in  the 
southern  side  of  Massachusetts  Bay  near  Plym- 
outh.41 Thus  stray  individuals  may  be  expected 
to  visit  the  southern  part  of  the  Gulf  in  most 
summers,  though  we  have  never  met  it  there 
ourselves.  It  has  even  been  reported  as  far  north 
as  Seguin  Island,  Maine,  but  without  convincing 
evidence  that  the  shark  in  question  was  not  a 
porbeagle.42 

Importance. — The  chief  importance  of  the  At- 
lantic mako,  as  of  its  Indo-Pacific  relative,  is  as  a 
game  fish,  because  of  its  fast  runs  when  hooked 
and  of  its  habit  of  leaping.  But  it  is  not  plentiful 
enough  anywhere  in  the  Gulf  of  Maine  to  be 
worth  fishing  for  there  especially. 

Maneater  Carcharodon  carcharias  (Linnaeus)  1758 
White  shark 

Bigelow  and  Schroeder,  1948,  p.  134. 
Garman,  1913,  PI.  5,  figs.  5-9. 

Description. — The  maneater  is  of  the  general 
"mackerel  shark"  appearance,  with  firm  lunate 
tail,  the  upper  lobe  only  a  little  longer  than  the 
lower ;  and  with  triangular  first  dorsal  of  moderate 
size  originating  over  the  armpits  of  the  pectorals, 
which  are  sickle  shaped,  and  roughly  twice  as 
long  as  they  are  broad.  The  second  dorsal  and 
anal  fins  are  very  small,  the  former  a  little  in 
advance  of  the  latter;  and  the  root  of  the  tail 

*'  Informal  ion  from  Dr.  W.  J.  Mixter. 

41  Various  early  reports  of  It  in  the  northern  part  of  the  Gulf  seem  to  have 
referred,  actually,  to  the  porbeagle. 


bears  a  single  well-marked  keel  on  either  side. 
The  snout  is  conical,  moderately  pointed. 

Unfortunately,  there  is  no  obvious  field  mark 
to  distinguish  a  small  maneater  from  a  large 
porbeagle  or  from  a  large  mako  when  seen  swim- 
ming at  any  distance.  Once  captured,  however, 
no  confusion  could  arise,  for  instead  of  the  slim 
catlike  teeth  of  the  porbeagle  and  of  the  mako,  we 
find  the  maneater  one  of  the  best  armed  of  all 
sharks ;  its  teeth  large  and  triangular,  and  similar 
in  shape  in  the  two  jaws,  except  broadest  in  the 
upper,  with  nearly  straight  cutting  edges  and 
strongly  serrated  margins.  As  a  precaution,  any 
large  active  shark,  upwards  of  10  or  12  feet  long, 
with  the  tad  not  long,  out  of  ordinary  proportions, 
should  be  looked  upon  with  suspicion,  for  it  might 
prove  to  be  a  maneater.  If  it  were  sluggish, 
resting  with  the  dorsal  fin  high  out  of  water,  it 
would  be  no  doubt  a  harmless  basking  shark 
(p.  28). 

Color. — Maneaters  up  to  12  to  15  feet  long  are 
slaty  brown  or  leaden  gray  above,  sometimes 
almost  black,  shading  more  or  less  abruptly  on  the 
sides  to  dirty  white  below.  There  is  a  black  spot 
in  the  armpit  of  each  pectoral  fin,  and  the  lower 
surfaces  of  the  pectorals  are  black  toward  their 
tips,  usually  with  some  black  spots  adjacent.  The 
pelvics  are  white  below,  but  olive  along  their 
anterior  edges.  Larger  specimens  (we  have  seen 
none)  have  been  described  as  dun  colored  above 
or  very  pale  leaden,  and  they  may  lack  the  black 
spot  at  the  armpit  of  the  pectoral  fin.43 

1 1nformation  from  Stewart  Springer,  from  large  Florida  specimens. 


Figure  7. — Maneater  (Carcharodon  carcharias),  Massachusetts,  about  7  feet  long.     A,  first  three  upper  and  B,  first 
three  lower  teeth,  from  center  of  jaw,  from  a  specimen  about  8%  feet  long,  Woods  Hole,  about  0.6  times  natural 
size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 
210941—53 3 


26 


FISHERY   BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


Size.- — This  is  one  of  the  largest  of  sharks.  A 
gulf  of  Maine  specimen  about  3  feet  long  is  the 
smallest,  apart  from  embryos,  that  has  been  seen; 
one  of  about  5  feet  the  next  smallest.  So  far  as 
known  it  does  not  mature  sexually  until  it  has 
grown  to  a  length  of  12  to  14  feet.  Among  larger 
ones,  from  one  place  or  another,  the  exact  measure- 
ments for  which  have  been  reported,  four  have 
been  between  14  and  16  feet  long,  two  between  16 
and  18  feet,  and  three  between  19  and  21  feet. 
The  largest  on  record  was  36%  feet  long;  **  the  next 
largest  about  30  feet,  but  perhaps  not  measured 
exactly. 

Maneaters  of  a  given  length  may  vary  widely 
in  weight,  because  of  variations  in  their  condition. 
Thus  one  specimen  8  feet  2  inches  long  weighed 
only  342  pounds,  but  another  of  8  feet  3  inches, 
weighed  600  pounds.  Five,  weighing  between  910 
and  1,000  pounds  ranged  from  9  feet  8  inches  in 
length  to  12  feet  6  inches.  Three,  of  13  to  13% 
feet,  weighed  1,291  to  1,344  pounds,  but  another, 
from  South  Africa  of  13  feet  3  inches  scaled  2,176 
pounds,  doubtless  a  very  fat  fish.  A  15-foot 
2-inch  specimen  weighed  1 ,720  pounds ;  and  one  of 
21  feet,  the  largest  that  has  been  weighed  so  far, 
7,100  pounds,  its  liver  1,005  pounds.46 

Habits. — So  few  maneaters  are  seen  that  little  is 
known  of  their  way  of  life,  apart  from  their  vorac- 
ity. Most  of  the  records  of  them  have  been  of 
specimens  taken  at  or  near  the  surface,  and  such 
specimens  as  visit  our  Gulf  sometimes  come  very 
close  inshore.  Thus  two  specimens  were  seined 
close  in,  off  Swampscott,  at  the  northern  entrance 
to  Boston  Harbor  in  1939;  one  was  harpooned  in 
1937  about  2  miles  off  Nantasket  Beach,  one  of 
the  most  popular  bathing  resorts  near  Boston; 
another  was  harpooned  about  one-half  mile  off 
Cohasset,  Mass.,  where  the  water  is  not  over  20 
feet  deep;  one  in  10  feet  of  water  in  Provincetown 
Harbor,  many  years  ago.  Some  have  even  been 
taken  in  fish  traps  close  to  the  beach  on  Cape  Cod 
and  near  Woods  Hole;  and  in  1916  one  was  taken 
in  the  shallow  water  of  Sandy  Hook  Bay,  N.  Y. 
On  the  other  hand,  the  largest  one  that  has  been 
weighed  yet  was  caught  on  a  set  line  off  the  north 
coast  of  Cuba,  at  a  depth  of  about  700  fathoms. 

Nothing  is  known  of  its  breeding  habits,  beyond 


«  This  Australian  specimen,  the  jaws  of  which  are  In  the  British  Museum, 
is  the  basis  for  repeated  statements  that  the  maneater  grows  to  40  feet. 

11  For  further  details,  see  BIgelow  and  Schroeder,  Fishes  Western  North 
Atlantic.  Pt.  1, 1948,  pp.  137-138. 


the  bare  facts  that  it  is  ovoviviparous  like  others 
of  the  mackerel  shark  tribe. 

The  maneater  is  one  of  the  most  voracious  of  all 
the  fish  tribe,  feeding  indifferently  on  large  prey 
and  on  small.  Other  sharks,  4  to  7  feet  long  and 
practically  intact,  have  been  found  repeatedly  in 
maneaters'  stomachs;  and  a  young  sea  lion  of  100 
pounds  in  one  on  the  coast  of  California,  while 
seals,  sturgeons,  and  tuna  have  been  found  in 
maneaters  no  longer  than  8  to  9  feet.  In  southern 
seas  they  are  described  as  feeding  regularly  on  sea 
turtles.  But  they  also  devour  smaller  fishes  of 
whatever  kinds  are  available,  including  small 
sharks  and  chimaeroids,  also  squids.  When  they 
come  in  on  the  fishing  banks,  they  are  known  to 
take  fish  that  they  find  hooked  on  long  lines  as 
porbeagles  do  (p.  22).  Thus  the  mouth  of  one  of 
9  feet  8  inches,  taken  near  Cohasset,  Mass.,  and 
examined  by  us,  carried  several  hooks  with  the 
snoods  still  attached,  while  its  stomach  contained 
a  spiny  dogfish  (Squalus  acanthias)  that  evidently 
had  been  torn  off  a  hook.  And  a  large  Florida 
maneater,  caught  on  a  set  line,  contained  2  brown 
sharks  (Carcharhinus  milberti) ,  6  to  7  feet  long,  that 
had  evidently  been  torn  from  hooks  on  the  same 
set  line  on  which  the  maneater  was  hooked.  The 
maneater,  like  the  Tiger  shark,  is  not  above  feed- 
ing on  slaughterhouse  waste  or  other  garbage. 

General  range. — This  is  an  oceanic  shark, 
widespread  in  the  tropical  and  warm  temperate 
belts  of  all  oceans,  including  the  Mediterranean. 
In  the  western  side  of  the  Atlantic  it  has  been 
recorded  as  far  north  as  St.  Pierre  Bank  south 
of  Newfoundland,  and  as  far  south  as  Brazil.46 

Occurrence  in  the  Gulf  of  Maine. — The  maneater 
is  usually  looked  on  as  a  warm  water  shark, 
doubtless  correctly  so.  None  the  less,  it  has  been 
reliably  reported  from  the  southwestern  part  of 
the  Gulf  of  Maine  more  often  than  it  has  from  any 
other  coastal  sector  of  comparable  length  on  the 
Atlantic  coast  of  North  America.  At  least  10, 
for  example,  were  actually  captured  or  were 
harpooned  and  lost  in  Massachusetts  Bay  alone 
during  the  period  1935  to  1948.  We  ourselves 
examined  tbree  of  these,  one  that  was  netted  at 
Swampscott;  a  female  of  9  feet  8  inches  weighing 
980  pounds  that  was  harpooned  within  half  a  mile 
of  the  land  off  Cohassett,  in  August  1940;  one  of 
about  3  feet,  that  was  harpooned  in  July  1948 

"  For  details  and  references,  see  Blgelow  and  Schroeder,  Fishes  Western 
North  Atlantic.  Pt.  1,  1948,  pp.  140-141. 


FISHES   OF  THE  GULF   OF   MAINE 


27 


near  Boston  Lightship,  this  last  being  the  smallest 
that  is  on  record  to  date  (p.  26),  and  one  about  14 
feet  long,  weighing  1,050  pounds  dressed,  which 
sold  for  10  cents  a  pound,  was  taken  in  a  trap  at 
North  Truro  on  November  9,  1952. 

Carrying  the  record  back  to  earlier  years,  a 
15-foot  shark,  taken  at  Monomoy  Point  at  the 
elbow  of  Cape  Cod  in  the  autumn  of  1928,  appears 
to  have  been  a  maneater,  and  one  of  about  16 
feet,  taken  in  a  trap  at  East  Brewster,  October  16, 
1923,  and  identified  by  Dr.  Samuel  Garman, 
certainly  was,  while  one  of  7  feet  2  inches,  taken 
in  Massachusetts  Bay,  about  1910,  was  the  basis 
of  Garman's  (1913,  pi.  5,  fig.  5)  beautiful  illus- 
tration. Earlier  still,  a  13-footer,  taken  at 
Provincetown,  Cape  Cod,  in  June  1848,  was 
described  by  Storer  as  a  new  species,  atwoodi, 
while  two  small  ones  were  mentioned  by  him  as 
taken  by  Massachusetts  fishermen  between  1820 
and  1850.  And  Capt.  Atwood  reported  seeing 
four,  caught  in  mackerel  nets  at  Provincetown 
many  years  ago.47 

Proceeding  northward,  we  find  scattered  records 
from  the  vicinity  of  Portland,  Maine,  most  re- 
cently, a  13-footer  caught  in  a  gill  net  off  Casco 
Bay  in  November  1931;  one  from  Eastport, 
Maine,  many  years  ago;  a  very  large  one  (esti- 
mated as  about  26  feet  long)  taken  in  a  wier  at 
Campobello  Island,  November  23,  1932  *8  it  was 
suggested  locally  that  it  may  have  been  the  same 
specimen  that  had  attacked  a  fishing  boat  off 
Digby,  Nova  Scotia,  the  preceding  July  (p.  27); 
one  from  Deer  Island,  New  Brunswick,  taken  in  a 
herring  weir,  August  24,  1949  ;49  and  one  from 
Digby,  on  the  Nova  Scotian  shore  of  the  Bay  of 
Fundy,  July  2,  1932.  And  there  are  several  re- 
liable records  for  St.  Margaret  Bay  on  the  outer 
coast  of  Nova  Scotia,  perhaps  also  for  Halifax. 

The  most  northerly  positive  record  for  it  on  the 
Atlantic  coast  of  North  America  is  for  St.  Pierre 
Bank,  south  of  Newfoundland,  where  one  attacked 
a  fisherman  in  a  dory  many  years  ago,  leaving 
in  the  sides  of  the  boat  pieces  of  its  teeth,  from 
which   Dr.    Garman   was   able   to   identify   it.50 

Westward  and  southward  from  the  elbow  of 
Cape  Cod,  we  find  nine  or  ten  definite  records  for 
Nantucket  and  for  the  vicinity  of  Woods  Hole 

«  Putnam.  Bull.  Essex  Inst.,  vol.  6,  1874,  p.  72. 
•'  Piers,  Proc.  Nova  Scotian  Inst.  Set.,  vol.  18, 1934,  p.  198. 
*  A  female  12  feet ,  8  Inches  long,  weighing  1 ,299  pounds,  reported  by  Scatter- 
good,  Trefetben,  and  Coffin,  Copela,  1961,  p.  298. 
»  Putnam,  Bull.  Essex  Inst.,  Salem,  vol.  6, 1874,  p.  72. 


(never  more  than  two  in  any  one  year),  with  one 
of  five  feet  (second  smallest  on  record)  netted  at 
Sakonnet,  Rhode  Island,  May  30,  1939.  Maneat- 
ers  are  also  reported  occasionally  near  New  York, 
notably  one  of  about  seven  feet,  taken  in  Sandy 
Hook  Bay,  July  1916,  to  which  we  recur  below 
(p.  27). 

Belation  to  man. — So  few  man-eaters  visit  our 
Gulf  that  they  would  deserve  only  the  briefest 
mention  were  this  not  the  only  shark  that  is  ever 
likely  to  attack  human  beings  there.  Strong  and 
active,  equipped  as  it  is  with  a  most  terribly 
effective  set  of  cutting  teeth,  it  has  borne  an  un- 
savory reputation  as  a  man-eater  from  the  earliest 
times,  and  it  is  probable  that  the  7-foot  specimen 
listed  earlier  from  South  Amboy,  Sandy  Hook 
Bay,  was  the  cause  of  the  shark  fatalities  along 
the  New  Jersey  beach  in  July  1916  (p.  16).  A 
fatal  attack  on  a  swimmer  at  Mattapoisett,  on 
Buzzards  Bay,  on  July  25,  1936,  may  also  have 
been  by  a  man-eater,  though  in  this  case  the 
shark  was  driven  away  without  being  identified. 

This  is  also  perhaps  the  only  shark  against  which 
unprovoked  attacks  on  small  boats  are  proved 
by  identification  of  their  teeth,  embedded  in  the 
wood.  One  such  instance,  from  the  Newfound- 
land Banks,  was  reported  by  Putnam "  many 
years  ago  (p.  27).  A  recent  local  case  is  of  a  very 
large  one  that  attacked  a  fishing  boat  in  the  Bay 
of  Fundy  off  Digby  Gut,  Nova  Scotia,  July  2, 
1932  and  left  in  her  keel  or  lower  planking  several 
of  its  teeth,  by  which  it  was  identified.62  Storer  63 
wrote  of  a  case  where  one  (apparently  the  13-foot 
specimen  that  he  had  described  earlier  as  atwoodi) 
turned  furiously  on  a  boat,  but  was  lanced  to  death 
and  brought  into  Provincetown.  And  a  15-foot 
shark,  probably  this  species  to  judge  from  the  il- 
lustration of  it  that  was  published,64  that  was 
killed  off  Monomoy  Point  by  two  fishermen  in 
November  1928,  overturned  their  dory  before  it 
was  subdued.  And  one  of  about  15  feet  (similarly 
identified  by  teeth  left  in  the  planking)  attacked 
a  boat,  from  which  it  had  been  harpooned,  in  St. 
Margaret's  Bay,  Nova  Scotia,  on  June  27,  1920." 
Hence,    so    long   as   maneaters    wander    within 


»'  Proc.  Essei  Inst.  Salem,  vol.  6,  18/4.  p.  72;  teeth  Identified  by  Dr.  S. 
Garman. 

"  Reported  by  Piers,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  18, 1934,  p.  198. 

«  Fishes  of  Mass.,  1867,  p.  248. 

*•  Reported  In  Wltman  and  Lee  Co.'s  Market  Letter  for  Nov.  8, 1928;  called 
to  our  attention  by  Dr.  Lewis  Radcllfle  of  the  U.  S.  Bureau  of  Fisheries. 

*»  For  details  of  this  occurrence,  see  Piers,  Proc.  Nova  Scoila  Inst.  Scl.,  vol. 
18,  1934,  pp.  196-198. 


28 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


our  limits  more  often  than  had  been  realized  pre- 
viously, the  possibility  is  always  open  of  attacks 
on  bathers  along  the  Massachusetts  shores  of 
the  Gulf. 

Despite  its  ferocity,  muscular  strength  and  size, 
the  man-eater  does  not  put  up  so  spectacular  a  re- 
sistance when  hooked  as  does  a  mako,  neither 
running  so  fast  nor  having  the  habit  of  jumping. 
Neither  does  it  put  up  as  strong  a  fight,  pound 


for  pound,  as  a  tuna  ordinarily  does,  or  any  of  the 
swordfish  tribe.  Thus  a  1,329-pound  maneater 
was  landed  on  rod  and  reel  by  an  Australian  angler 
in  53  minutes.  One  of  2,176  pounds,  caught 
from  the  shore  in  South  Africa,  is  the  largest  fish 
ever  landed  on  rod  and  reel  that  has  come  to  our 
notice.66 


»  London  Illus.  News,  July  14, 1928,  p.  63;  photograph  recorded  as  a  mako 
but  shown  by  its  teeth  to  have  been  a  maneater. 


BASKING  SHARKS.  FAMILY  CETORHINIDAE 


Basking  shark  Cetorhinus  maximus   (Gunnerus) 
1765 

Bone  shark 

Bigelow  and  Schroeder,  1948,  p.  147. 

The  basking  shark  resembles  the  mackerel 
sharks  in  the  lunate  shape  of  its  caudal  fin,  with 
lower  lobe  nearly  as  long  as  upper;  also  in  the 
presence  of  a  noticeable  lunate  furrow  above  and 
one  below  on  the  root  of  the  tail,  and  in  the 
wide  lateral  expansion  of  the  latter,  forming  a  pro- 
nounced "fore  and  aft"  keel  on  either  side;  also 
in  the  facts  that  the  second  dorsal  fin  and  the  anal 
fin  are  much  smaller  than  the  first  dorsal,  that  its 
fifth  gill  opening  is  situated  in  front  of  the  origin 
of  the  pectoral  fin;  in  the  position  of  the  mouth  on 
the  under  side  of  the  head ;  and  in  the  wide  separa- 
tion of  the  nostrils  from  the  mouth.     But  the  teeth 


of  the  basking  shark  are  minute  and  very  numer- 
ous (large  and  few  in  number  in  the  mackerel 
sharks);  its  gill  openings  are  so  large  that  they 
extend  right  around  the  neck,  with  those  of  the 
first  pair  almost  meeting  below  on  the  throat ;  and 
the  inner  margin  of  each  gill  arch  bears  a  great 
number  of  horny,  bristle-like  rakers,  directed 
inward-forward,  that  correspond  to  the  rakers  of 
various  bony  fishes  in  their  position  and  in  their 
function  (see  p.  30).  It  was  the  fancied  resem- 
blance of  these  rakers  to  the  whalebone  of  the 
whalebone  whales  that  suggested  the  vernacular 
name  "bone  shark"  to  the  whalemen  of  olden 
times. 

Corresponding  to  its  feeding  habits,  the  mouth 
of  the  basking  shark  is  very  large  and  widely  dis- 
tensible at  the  corners.  The  snout  is  short, 
conical,  with  rounded  tip  on  large  specimens. 
But  it  is  much  longer,  relatively,  on  small  ones, 


Figure  8. — Basking  shark  (Cetorhinus  maximus),  26J4-foot  female,  Marthas  Vineyard.  A,  side  view  of  head  of  12-foot 
Long  Island  specimen;  B,  a  group  of  the  teeth  of  same,  about  1.2  times  natural  size.  From  Bigelow  and  Schroeder. 
Drawings  by  E.  N.  Fischer. 


FISHES   OF  THE  GULF  OF   MAINE 


29 


projecting  far  beyond  the  mouth,  obliquely  trun- 
cate in  front,  terminating  above  in  a  sharp  point, 
and  with  the  head  strongly  compressed  sideways 
abreast  of  the  front  of  the  mouth.  This  results  in 
so  bizarre  an  appearance  that  the  young  basking 
shark  was  thought  at  first  to  represent  a  separate 
species.  A  gradual  transition  takes  place  from 
the  juvenile  shape  of  head  to  the  adult  shape  when 
a  length  of  12  to  16  feet  has  been  reached.  We 
need  only  note  further  that  the  triangular  first 
dorsal  fin  stands  midway  between  pectorals  and 
pelvics;  though  not  so  high  in  proportion  as  that 
of  the  mackerel-shark  tribe,  it  rises  high  in  the  air 
when  a  large  basking  sbark  lies  awash  on  the  sur- 
face, as  is  their  habit,  a  convenient  field  mark 
(p.  29). 

Color. — Upper  surface  grayish  brown,  slaty 
gray,  or  even  almost  black.  The  lower  surface 
has  been  described  repeatedly  as  white.  But  the 
Menemsha  specimen  described  by  Allen  67  was  of  a 
somewhat  lighter  shade  below  than  above,  without 
white  markings,  as  was  a  Massachusetts  Bay 
specimen  recently  examined  by  us;  while  one  14 
feet  long  captured  at  West  Hampton,  L.  I., 
on  June  29,  1915  68  had  the  belly  as  dark  as  the 
back,  with  a  white  patch  underneath  the  snout  in 
front  of  the  mouth. 

Size. — The  basking  shark  rivals,  though  it  does 
not  equal,  the  whale  shark  of  tropical  seas  in  size. 
Reports  that  an  occasional  basking  shark  may 
reach  a  length  of  50  feet  probably  are  not  an 
exaggeration,  for  the  catch  on  the  coast  of  Norway, 
for  the  period  1884  to  1905,  included  one  of  about 
45  feet  and  three  of  about  40  feet,  with  the  six 
next  longest  ranging  between  36  feet  and  30  feet  3 
inches.  The  three  longest  for  which  we  find 
definite  measurements  for  the  western  Atlantic 
were  of  32  feet  2  inches,  32  feet,  and  30  feet  3 
inches.  But  others  up  to  35  feet  long  have  been 
credibly  reported  as  killed  near  Eastport,  Maine, 
many  years  ago;  and  one  captured  at  Musquash 
Harbor,  New  Brunswick,  near  the  mouth  of  the 
Bay  of  Fundy  in  1851  was  said  to  have  been  about 
40  feet  long.  It  is  probable  that  they  are  at  least 
5  to  6  feet  long  when  born,  the  three  smallest  so  far 
reported  having  been  between  5  feet  5  inches  and 
about  8  feet  6  inches  long.  Matthews  59  concluded 
from  studies  of  basking  sharks  taken  near  the  Isle 

>'  Bull.  Boston  Soc.  Nat.  Hist.,  No.  24,  March  1921,  p.  5. 

"  Described  by  Hussakof,  Copela,  No.  21, 1915,  pp.  25-27. 

«  Philos.  Trans.  Roy.  Soc.  London,  Ser.  B.,  vol.  234, 1950,  pp.  217-316. 


of  Skj'e  that  fish  up  to  10  feet  are  in  their  first 
year,  those  of  15  feet  in  their  second  year.  Males 
mature  sexually  at  about  18  to  20  feet  as  indicated 
by  the  lengths  of  their  claspers,  females  at  about  20 
to  23  feet;  i.  e.,  when  3  years  old  or  perhaps  4, 
according  to  Mathews'  estimate. 

We  find  no  exact  weights  for  large  basking 
sharks  from  the  Atlantic.  But  6,580  pounds  for 
one  of  28  feet,  and  8,600  pounds  for  another  of  30 
feet,  from  Monterey,  Calif.,  is  doubtless  a  fair 
indication  of  what  a  fairly  large  one  may  be 
expected  to  weigh.  Estimated  weights  for  smaller 
ones,  from  the  Pacific,  are  about  6,600  pounds  at 
about  23  feet,  1,000  to  1,800  pounds  at  13  to  15 
feet,  and  800  pounds  at  8  feet  4  inches.60  A  young 
one,  12  feet  long,  killed  off  Digby,  Nova  Scotia, 
August  16,  1939,  weighed  359  pounds,  after  it  had 
bled, 61  and  one  almost  20  feet  long,  taken  off 
Portland,  Maine,  in  1936,  weighed  550  pounds, 
dressed. 

Habits. — This  is  a  sluggish,  inoffensive  fish,  help- 
less of  attack  so  far  as  its  minute  teeth  are  con- 
cerned. It  spends  much  time  sunning  itself  at 
the  surface  of  the  water,  often  lying  with  its  back 
awash  and  dorsal  fin  high  out  of  water,  or  on  its 
side,  or  even  on  its  back  sunning  its  belly;  some- 
times it  loafs  along  with  the  snout  out  of  water, 
the  mouth  open,  gathering  its  provender  of  plank- 
ton. They  pay  so  little  attention  to  boats  that 
it  is  easy  to  approach  one  of  them  within  harpoon 
range,  and  excellent  motion  pictures  have  beeo. 
taken  of  them  in  Irish  waters.62  But  they  have 
also  been  seen  jumping,  perhaps  to  shake  off  para- 
sites. Those  seen  in  the  Gulf  of  Maine  are  usually 
traveling  singly.  But  they  are  known  to  congre- 
gate sometimes  in  loose  schools  which  may  include 
as  many  as  60  to  100  in  the  peak  years  of  abun- 
dance for  them  in  regions  where  they  are  more 
numerous  than  in  the  Gulf  of  Maine.63  It  is 
chiefly  during  the  warm  half  of  the  year  that 
basking  sharks  are  encountered  off  the  northeast- 
ern United  States  and  in  the  northern  part  of  then- 
range  in  the  opposite  side  of  the  Atlantic.  It  is 
likely  that  those  that  summer  in  the  inshore  parts 
of  the  Gulf  simply  withdraw  in  the  fall,  to  pass  the 


M  For  further  details  as  to  sizes  of  basking  sharks,  see  Bigelow  and 
Schroeder,  Fishes,  Western  North  Atlantic,  Pt.  1, 1948,  pp.  151-152. 

"  Referred  to  by  McKenzie,  F-roc.  Nova  Scotia  Hist.  Sci..  vol.  20, 1940,  p.  42. 

"  Shown  In  the  film  "Men  of  Arran." 

u  See  Bigelow  and  Schroeder,  Fishes  Western  North  Atlantic,  Pt.  1, 
1948,  pp.  153,  154,  for  details  as  to  their  centers  of  population  and  seeular 
fluctuations  in  abundance  in  north  European  waters. 


30 


FISHERY   BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


winter  in  deeper  water  where  the  temperature 
does  not  fall  so  low. 

Next  to  its  vast  bulk  and  its  curiously  sluggish 
habit,  the  most  interesting  peculiarity  of  the  bask- 
ing shark  is  its  diet,  for  it  subsists  wholly  on  tiny 
pelagic  animals,  which  it  sifts  out  of  the  water  by 
means  of  its  greatly  developed  gill  rakers,  exactly 
as  plankton-feeders  among  fishes  such  as  men- 
haden do,  and  whalebone  whales  with  their  baleen 
sieves.  In  several  cases  their  stomachs  have  been 
found  packed  with  minute  Crustacea;  this  was  true 
of  the  only  western  Atlantic  specimen  of  which 
the  stomach  contents  have  been  examined.  And 
while  digestion  is  so  rapid  that  the  food  swallowed 
is  soon  reduced  to  a  soupy  mass,  this  usually  is 
reddish,  suggesting  a  crustacean  origin. 

All  that  is  known  of  the  breeding  of  the  basking 
shark  is  that  the  structure  of  the  internal  sex  or- 
gans of  the  female  accords  with  the  nourishment 
of  the  embryo  within  the  maternal  oviduct, 
that  the  ovary  of  a  female,  with  empty  oviduct 
contained  something  like  6  million  immature  ova 
instead  of  the  few  that  are  usual  in  sharks  that 
bear  "living"  young,  and  that  an  embryo  about  a 
foot  long  was  said,  long  ago,  to  have  been  taken 
from  its  mother.64 

Basking  sharks  reported  as  "sea  serpents"  or  as 
other  "monsters" . — The  remains  of  basking  sharks 
have  been  reported  as  "sea  serpents"  on  several 
occasions;  nor  is  this  astonishing.  "As  the  carcass 
of  the  shark  rots  on  the  shore,  or  is  buffeted 
against  the  rocks,  the  whole  of  the  gristly  skeleton 
of  the  jaws  and  gill  arches  ...  as  well  as  the 
pectoral  and  pelvic  fins,  is  soon  washed  away,"65 
leaving  only  the  cranium  and  the  long  backbone, 
with  larger  or  smaller  amounts  of  muscle,  so  frayed 
out  as  to  suggest  a  hairy  or  bristly  mane.  As 
a  recent  instance  from  the  Gulf  of  Maine,  we  may 
cite  the  newspaper  and  radio  publicity,  that  was 
given,  as  a  supposed  sea  serpent,  to  a  basking 
shark  skeleton,  about  25  feet  long,  that  beached 
near  Provincetown  on  the  outer  shore  of  Cape 
Cod,  in  January  1937,  that  we  examined.66 

A  more  spectacular  instance  of  the  fanciful  in- 
terpretation that  is  likely  to  be  placed  on  any 
large  stranded  carcass  that  has  decayed  partially, 
was  the  famous  "Animal  of  Stronsa,"  that  came 

«  See  Matthews,  Philos.  Trans.  Eoy.  Soc.  London,  Ser.  B,  No.  612,  vol. 
234, 1960,  pp.  347—366  for  detailed  account. 

"  Norman  and  Fraser.  Giant  Fishes,  Whales  and  Dolphins,  1937,  p.  21. 

*  For  account  and  photograph,  see  Schroeder,  New  England  Naturalist, 
No.  2, 1939,  p.  1. 


ashore  on  the  island  of  that  name  in  the  Orkneys, 
in  September  1808.  It  was  pictured  by  an  eye- 
witness as  having  three  pairs  of  limbs,  but  the 
published  illustration  of  its  cranium,  vertebrae, 
and  pelvic  skeleton  6:  show  that  it  was  only  the 
remains  of  some  very  large  shark,  probably  a 
basking  shark.  It  has  also  been  suggested  repeat- 
edly that  some  of  the  stories  of  sea  monsters  of 
one  sort  or  another  may  have  been  based  on  the 
dorsal  and  caudal  fins  of  two  or  more  basking 
sharks,  swimming  one  behind  another  as  they 
often  do  (we  dare  not  touch  further  on  the  contro- 
versial subject  of  the  "sea  serpent"). 

General  range. — This  enormous  fish,  formerly 
thought  to  be  an  Arctic  species,  straying  south- 
ward, is  now  known  to  be  an  inhabitant  of  the 
temperate-boreal  zone  of  the  North  Atlantic.68 
It  is  represented  in  the  corresponding  thermal 
belts  of  the  South  Atlantic  and  of  the  North 
and  South  Pacific  by  a  similar  great  shark  (or 
sharks),  whose  exact  relationship  to  the  basking 
shark  of  the  North  Atlantic  is  still  an  open  question. 

The  northern  boundary  of  the  normal  range 
of  the  basking  shark  of  the  North  Atlantic  appears 
to  follow  the  line  of  transition  from  waters  of 
predominately  Atlantic  influence  to  those  of 
Arctic  origin.  This,  roughly,  runs  from  the  outer 
coast  of  Nova  Scotia  (1  record),  and  from  southern 
Newfoundland  (4  positive  records)  to  western  and 
southern  Iceland,  to  the  Orkney  and  Faroe  Islands, 
and  skirts  the  Norwegian  coast  to  the  North 
Cape,  while  basking  sharks  stray  now  and  then 
to  the  Murman  coast.  To  the  southward,  in  the 
North  Atlantic,  they  range  as  far  as  the  Mediter- 
ranean and  Morocco  in  the  east,  to  North  Carolina 
in  the  west. 

Occurrence  in  the  Gulf  of  Maine. — Before  the 
coming  of  the  white  man  this  great  shark  seems 
to  have  been  a  regular  inhabitant  of  the  southern 
part  of  the  Gulf  of  Maine.  And  tradition  has 
it  that  large  numbers  were  taken  in  Massachusetts 
waters,  especially  off  the  tip  of  Cape  Cod,  during 
the  first  half  of  the  eighteenth  century,  for  their 
liver  oil  which  was  then  in  demand  for  illuminating 
purposes.  However,  the  local  stock  seems  soon 
to  have  gone  the  same  way  as  the  local  stock  of 
the  North  Atlantic  right  whale;  that  is,  into  the 
try  pot.     And  basking  sharks  seem  never  to  have 

i  Barclay,  Mem.  Wernerlan  Soc,  Edinburgh,  vol.  1, 1811,  p.  418. 

•!  It  has  long  been  realized  that  old  tales  of  a  tremendous  whale-eating 
shark,  on  which  Fabricius  based  his  statement  that  the  basking  shark  occurs 
In  Greenland  waters,  were  fiction. 


FISHES   OF  THE  GULF  OF   MAINE 


31 


visited  the  northeastern  part  of  the  Gulf  in  any 
numbers,  there  being  only  a  few  records  for  the 
vicinity  of  Eastport,  Maine,  and  three  from  within 
the  Bay  of  Fundy.  At  the  present  time  the  Gulf 
appears  to  harbor  a  sparse  and  fluctuating  popu- 
lation, occasional  members  of  which  are  encoun- 
tered from  time  to  time,  here  or  there,  but  whether 
as  immigrants  into  the  Gulf  from  the  open  ocean 
is  not  known. 

The  list  of  specimens,  the  capture  or  stranding 
of  which  in  the  Gulf  has  come  to  our  attention 
for  the  period  1908-1951  is  as  follows: 

1908.  One,  18  feet  long,  near  Provincetown,  taken  in  a 
fish  trap;  measured  by  J.  Henry  Blake. 

1909.  One,  about  22  feet,  in  Provincetown  Harbor. 
1913.     One,  about  29  feet,  Provincetown. 

1925.     One,  about  29  feet,  near  Monhegan  Island,  Maine. 
1931.     Female,  12H  feet  long,  York  Harbor,  Maine. 
1934.     One,  29  feet,  Whale  Cove,  Grand  Manan  Island, 

and  one,  28  feet,  Back  Bay,  Bay  of  Fundy.69 
1936.     Two  off  Portland,  Maine;  the  first  about  20  feet 

long,   weighing  550  pounds  dressed,   about   May   1; 

the  second,  much  larger  (reported  as  of  about  40  ft.), 

August  2. 
1939.     Skeleton  of  one  of  about  25  feet,  examined  by  us, 

found  on  the  beach  near  Provincetown  in  January. 

One  of  about  25  feet,  Yarmouth,  Nova  Scotia.     One 

of  12  feet,  Bay  of  Fundy  off  Digby  Gut.70 
1947.     Female,  about  13  feet  long,  examined  by  us,  har- 
pooned by  W.  T.  Reid  3rd,  near  Boston  Lightship, 

August  5th. 
1949.     A  small  one   (size  not  recorded),  near  Rockport, 

Mass.,  September;  identified  from  a  good  photograph 

by    Miss   D.    E.    Snyder  of  the   Peabody    Museum, 

Salem. 
1951.     One,  12  feet,  near  Bar  Harbor,  Maine,  harpooned 

July  28." 

Occasional  basking  sharks  also  visit  the  shores 
of  the  southern  coast  of  Massachusetts,  westward 
from  Cape  Cod;  one,  for  example,  12  to  14  feet 
long  was  taken  at  Menemsha  on  Marthas  Vine- 
yard, August  16,  1916;  another  of  20  feet  6  inches 
at  that  same  locality  on  June  24,  1920;72  one 
20  feet  2  inches  long  was  stranded  in  Hadleys 
Harbor,  Naushon  Island,  July  1937;  and  one  of 
8  feet  (among  the  smallest  on  record)  was  taken 
in  a  fish  trap  near  Woods  Hole  on  June  15,  1948. 

M  McKenzle,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  20, 1939,  p.  14. 

"  McKenzle,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  20, 1939,  p.  14. 

71  Personal  communication  from  J.  W.  Burger. 

"  This  specimen,  mounted,  In  the  New  England  Museum  of  Science  and 
described  by  Allen  (Bull.,  Boston  Soc.  Nat.  Hist.,  No.  24,  March  1921,  pp. 
3-10),  served  as  chief  basis  for  the  illustration  given  here  of  the  adult  basking 
shark. 


Probably  the  basking  shark  is  no  more  plentiful 
near  shore  in  our  Gulf  in  most  years  than  the 
paucity  of  the  recent  records  suggest,  for  popular 
interest  in  sharks  is  now  so  keen,  as  represented  by 
newspaper  publicity  given  to  any  unusual  capture, 
that  any  well-grown  one  is  apt  to  be  seen  in  these 
frequented  and  hard-fished  waters.  We  do  not 
find  evidence  of  any  considerable  incursion  by 
them  into  coastal  waters  farther  west  since  1878, 
when  20,  at  least,  were  found  dead  in  the  fish  traps 
near  Woods  Hole  during  the  summer.  And  the 
only  report  that  might  be  based  on  the  basking 
shark  on  the  offshore  fishing  banks  that  we  have 
received  from  fishermen  has  been  of  a  number  of 
unusually  large  sharks  of  some  sort,  seen  by  Capt. 
Henry  Klimm  on  the  southeast  part  of  Georges 
Bank  during  late  June  and  early  July  1947. 

Importance. — The  day  of  any  regular  fishery  for 
the  basking  shark  is  long  since  past  in  New 
England  waters,  probably  never  to  return.  And 
no  use  is  made  there,  nowadays,  of  the  occasional 
specimens  that  are  captured.  But  it  may  be  of 
interest  to  point  out  that  it  was  always  hunted  of 
old  by  the  sperm  whalers  from  New  Bedford,  for 
its  liver  oil  was  considered  nearly  or  as  good  as 
sperm  oil  for  illuminating  purposes.  Basking 
sharks  are  still  the  object  of  intermittent  small 
vessel  fisheries  off  the  coast  of  Iceland,  around  the 
Orkneys,  off  western  Ireland,  and  off  southern 
Norway;  also  off  Ecuador  and  Peru  in  the  Pacific. 
And  increasing  numbers  have  been  landed  during 
the  past  few  years  in  northern  California,  where 
they  are  considerably  more  plentiful  than  they 
are  in  the  Gulf  of  Maine,73  for  fish  meal  and  for  the 
liver  oil.  The  yield  of  oil  per  fish  varies  from 
about  80  gallons  to  about  200,  occasionally  to  400 
gallons,  with  as  much  as  600  gallons  reported. 
The  liver  of  a  30-foot  fish  weighing  6,580  pounds, 
taken  off  Monterey,  Calif.,  had  a  liver  weighing 
1,800  pounds,  60  percent  of  which  was  oil.7*  But, 
sad  to  say,  it  is  very  low  in  vitamin  A. 

The  fishery,  wherever  carried  on,  is  by  harpoon. 
And  basking  sharks  are  so  sluggish  and  so  un- 
suspicious of  a  boat,  large  or  small,  that  it  usually 
is  a  simple  matter  to  harpoon  one  that  is  seen  at 


"According  to  MacGinitie  (Science,  N.  Ser.,  vol.  73,  1931,  p.  496),  21 
basking  sharks  were  landed  in  Monterey,  Calif.,  between  November  22, 1930 
and  February,  1931. 

'<  MacGinitie,  Science,  N.  Ser.,  vol.  73,  May  1931,  p.  496. 


32 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


the  surface.  Once  struck,  however,  a  large  one  is 
likely  to  put  up  an  astonishingly  active  and 
enduring  resistance.  We  read,  for  example,  of 
one  of  35  to  38  feet  harpooned  by  Capt.  N.  E. 


Atwood  off  Provincetown,  Mass.,  about  1863, 
that  towed  the  fishing  smack  all  night,  and  broke 
loose  finally.75 


'•  Ooode,  Fish.  Ind.  U.  S.,  1884,  Sect.  1,  p.  669. 


THRESHER  SHARKS.     FAMILY  ALOPIIDAE 


The  threshers  (several  species  are  known)  are 
peculiar  among  sharks  for  their  enormously  long 
tail  fin.  Their  closest  affinities  in  other  respects 
are  with  the  mackerel  sharks. 

Thresher  Alopias  vulpinus  (Bonnaterre)  1 758 

Thraser;  Swiveltail;  Fox  shark 

Bigelow  and  Schroeder,  1948,  p.  167. 
Garman,  1913,  pi.  7,  figs.  1-3. 

Description. — The  thresher  is  as  easily  distin- 
guished from  all  other  Gulf  of  Maine  sharks  by  its 
long  tail  as  the  hammerhead  is  by  its  head,  the 
upper  caudal  lobe  being  a  little  longer  than  the 
head  and  body  of  the  fish  together,  curved  much 
like  the  blade  of  an  ordinary  scythe,  and  notched 
near  the  tip,  whereas  the  lower  lobe  measured  along 
the  front  margin  is  hardly  longer  than  the  pelvic 
fins.  We  need  merely  point  out  in  addition  that 
the  first  dorsal  fin  (of  moderate  size  and  about  as 
high  as  it  is  long)  stands  about  midway  between 
pectoral  and  pelvic  fins;  that  the  second  dorsal 
fin  and  the  anal  are  very  small;  that  the  pectoral 
fin  is  long  and  sickle  shaped;  and  that  the 
thresher  is  a  stout-bodied  shark  with  short  snout 


and  blunt,  rounded  nose.  Its  teeth  are  small, 
subtriangular  with  a  single  sharp  cusp  and  are 
smooth  edged.  Those  near  the  center  of  mouth 
are  nearly  symmetrical,  but  the  successive  teeth 
are  increasingly  oblique  outward,  with  their  outer 
margins  increasingly  concave. 

Color. — Dark  brown,  blue-slate,  slate  gray,  blue 
gray,  leaden  or  even  nearly  black  above,  often 
with  metallic  luster,  grading  on  the  sides  to  white 
below,  except  that  the  snout  and  the  lower  surface 
of  the  pectorals  are  usually  about  as  dark  below 
as  above,  and  that  the  sides  near  the  pectorals 
may  be  more  or  less  mottled  with  gray,  the  belly 
also.     The  iris  is  black  or  green. 

Size. — Threshers  vary  considerably  in  size  at 
birth,  for  while  free  living  specimens  have  been 
reported  as  small  as  46  inches,  with  many  of  48  to 
60  inches  (some  with  umbilical  scars  still  showing), 
one  unborn  embryo  was  61  inches  long.  The  state 
of  development  of  the  claspers  of  males,  with  the 
lengths  (14  ft.  6  in.  and  about  15%  ft.)  of  females 
that  have  been  found  with  embryos,  makes  it 
unlikely  that  they  mature  sexually  until  they  are 
at  least  14  feet  long  (tail  included).     Lengths  up 


D^r^9 


^ 


\i 


Figure  9. — Thresher  (Alopias  vulpinus),  about  5  feet  long, 
Rhode  Island,  from  Goode,  drawing  by  H.  L.  Todd. 
A,  upper  second  tooth;  B,  upper  third  tooth;  C,  upper 
fifth  tooth;  D,  upper  fifteenth  tooth;  E,  lower  second 
tooth;  F,  lower  sixth  tooth,  counted  from  center  of 
jaw;  about  2  times  natural  size.  From  Bigelow  and 
Schroeder.     Drawings  by  E.  N.  Fischer. 


FISHES   OF  THE  GULF   OF   MAINE 


33 


to  16  feet  are  usual;78  the  maximum  length  (tail 
included)  is  about  20  feet.  Threshers  are  so  large- 
ly tail  that  they  are  much  lighter  than  many  other 
sharks,  length  for  length.  The  few  actually 
weighed  have  ranged  from  about  300  to  320 
pounds  at  about  10  feet,  and  375  to  400  pounds  at 
about  13  feet,  to  about  500  pounds  at  about  14% 
feet.  Perhaps  1,000  pounds  is  about  the  maxi- 
mum to  be  expected  for  a  very  large  one. 

Habits. — The  reports  of  threshers  are  mostly 
based  on  ones  seen  at  the  surface  or  caught  either 
in  nets  set  shoal,  or  in  traps  set  close  inshore.  But 
a  thresher  has  been  hooked  as  deep  as  35  fathoms 
in  British  waters.77 

The  thresher  feeds  chiefly  if  not  exclusively  on 
small  schooling  fishes;  in  American  waters  mostly 
on  mackerel,  menhaden,  herring,  and  bluefish 
(Pomalomus) ;  also  on  bonito  and  on  squid.  A 
pair  of  threshers  often  work  in  concert  "herding" 
a  school  of  fish,  and  it  is  to  frighten  its  prey  together 
that  the  enormously  long,  flail-like  tail  is  em- 
ployed. Allen  78  gives  an  interesting  eyewitness 
account  of  a  thresher  pursuing  and  striking  a 
single  small  fish  with  its  tail. 

The  tale  that  the  thresher  leagues  with  the 
swordfish  to  attack  whales  is  time  honored,  but 
has  long  since  been  relegated  to  the  category  of 
myth.  And  so  weak  toothed  is  this  shark  that  the 
second  part  of  the  story  (it  makes  a  meal  of  its 
huge  victim)  is  close  to  an  impossibility.  The 
thresher,  we  may  add,  does  not  harm  human 
beings. 

In  American  waters  it  is  probable  that  threshers 
are  born  throughout  its  range,  very  small  free 
living  specimens  having  been  caught  off  New  Eng- 
land on  the  one  hand,  and  off  Florida  on  the  other. 
The  embryos  do  not  develop  a  placental  attach- 
ment with  the  mother,  and  either  2  or  4  have  been 
reported  in  gravid  females. 

General  range. — This  is  an  oceanic  shark  of 
temperate  and  subtropical  seas.  In  the  Atlantic 
it  is  known  from  southern  Ireland  and  the  North 
Sea  to  Madeira  and  the  Mediterranean  in  the  east, 
and  also  from  the  Cape  of  Good  Hope ;  from  Nova 
Scotia  and  the  Gulf  of  St.  Lawrence  to  Cuba  and 
the  northern  part  of  the  Gulf  of  Mexico  in  the 

Tfl  Several  of  that  size  have  been  taken  in  the  traps  at  Woods  Hole. 

11  There  is  another  group  of  species  of  the  genus,  with  very  large  eyes,  that 
live  at  greiter  depths;  for  discussion  of  these,  see  Bigelow  and  Schroeder 
(Fish.  Western  North  Atlantic,  Pt.  1,  1948,  pp.  162, 163). 

"  Science.  N.  Ser.,  vol.  58, 19?3,  pp.  31-32. 


west,  and  again  from  southern  Brazil  and  northern 
Argentina.  Seemingly  it  does  not  occur  in  the 
equatorial  belt  of  the  Atlantic.  But  it  does  in  the 
Pacific,  where  it  is  known  from  Oregon  to  Panama 
and  Chile.  Threshers  of  this  same  type  are  also 
found  in  the  central  and  western  Pacific  and  in 
the  Indian  Ocean.  Whether  the  thresher  of  the 
eastern  side  of  the  Pacific  is  identical  with  that 
of  the  Indian  Ocean  remains  to  be  determined. 

Occurrence  in  the  Gulf  of  Maine. — The  thresher 
has  often  been  seen  off  the  southern  coast  of  New 
England  and  in  some  numbers.  Three  about  16 
feet  long  have  been  taken  near  Woods  Hole,  for 
example,  in  one  trap  in  a  single  morning,  and  it 
has  been  classed  as  the  commonest  of  the  large 
sharks  off  Block  Island.  Scattered  specimens 
also  visit  the  Gulf  of  Maine  in  some  years,  though 
perhaps  none  in  others.  Thus  two  have  been 
reported  in  print  from  Nantucket;  we  saw  several 
large  ones  in  Pollock  Rip,  off  the  southern  angle 
of  Cape  Cod  on  August  4,  1913;  it  has  been  re- 
ported repeatedly  on  the  coast  of  Massachusetts, 
as  at  Barnstable  on  Cape  Cod  Bay,  where  one 
about  10  feet  long  was  taken  in  a  trap  on  October 
21,  1949,  and  from  various  localities  in  Massa- 
chusetts Bay  (e.  g.  Boston  Harbor  and  Nahant). 

Records  for  it  along  the  coast  of  Maine  include 
the  vicinity  of  Monhegan  Island,  east  of  Matinicus 
Island,  the  offing  of  Penobscot  Bay  where  one 
weighing  about  500  pounds  (estimated)  was 
caught  in  1911  and  another  seen  in  1911,  in  the 
vicinity  of  Eastport.  It  has  also  been  taken  in 
the  cold  waters  of  Passamaquoddy  Bay;  one  for 
instance  in  a  weir  at  Deer  Island,  August  28, 
1936; 79  also  in  the  Basin  of  Minas  on  the  Nova 
Scotian  shore  of  the  Bay  of  Fundy.  Occasionally 
a  thresher  is  netted  or  seen  off  the  outer  coast  of 
Nova  Scotia.  The  most  northerly  record  for  it 
from  our  side  of  the  Atlantic  is  for  the  Bay  of 
Chaleur  in  the  southern  side  of  the  Gulf  of  St. 
Lawrence.  It  is  to  be  expected  in  Gulf  of  Maine 
waters  only  during  the  warm  half  of  the  year, 
perhaps  May  to  October  (April  to  late  autumn  for 
Woods  Hole);  in  the  cold  season  it  altogether 
deserts  our  northern  coasts  for  warmer  seas. 

Importance. — The  thresher  is  not  common 
enough  in  the  Gulf  of  Maine  to  be  of  any  impor- 
tance to  fishermen  one  way  or  another,  or  to  play 


'»  Reported  by  McKenzie,  Proc.  Nova  Scotia  Inst.  Sci.,  vol.  20, 1939,  p.  14. 


210941—53- 


34 


FISHERY   BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


a  practical  role  of  any  moment  among  the  smaller 
fish.  Further  south,  however,  and  wherever  it 
is  numerous  in  the  Atlantic,  it  makes  itself  a  pest, 


tangling  and  tearing  mackerel  nets  as  well  as 
destroying  and  chasing  away  the  more  valuable 
fishes  on  which  it  feeds. 


CAT  SHARKS.     FAMILY  SCYLIORHINIDAE 


Distinctive  features  of  these  little  sharks  are 
that  they  have  five  pairs  of  gill  openings  and  an 
anal  fin;  that  at  least  one-half  of  the  base  of  the 
first  dorsal  fin  is  rearward  of  the  point  of  origin 
of  the  pelvic  fins;  that  the  front  margin  of  the 
nostrils  does  not  bear  a  fleshy  barbel;  and  that 
they  lay  eggs  with  horny  shells  and  tendrils  at 
the  corners.  Many  species  are  known.  The 
familiar  spotted  dogfishes  of  European  seas  (two 
species)  fall  in  this  group.  And  one  species  calls 
for  mention  here. 

Chain  dogfish  Scyliorhinus  retifer  (Garman) 
1881 

Description. — The  chain-like  pattern  of  narrow 
black  stripes  with  which  the  reddish-brown  back 
and  sides  of  this  little  shark  are  marked  are  so 
distinctive  that  there  is  no  likelihood  of  confusing 
it  with  any  other  shark.  We  need  only  add  that 
its  first  dorsal  fin  stands  wholly  behind  the  rear 
ends  of  the  bases  of  its  pelvic  fins;  that  its  second 


dorsal  fin  is  about  one-half  as  large  in  area  as  its 
first  dorsal  fin;  that  its  tail  fin  is  square-tipped 
and  occupies  only  about  one-fifth  of  the  length 
of  the  fish;  and  that  its  teeth  are  similar  in  the 
two  jaws,  narrow-triangular  with  a  small  second- 
ary cusp  on  either  side. 

Size. — The  largest  specimen  measured  so  far 
was  17  inches  long. 

General  range  and  occurrence  in  the  Gulf  of 
Maine. — The  range  of  the  chain  dogfish  is  con- 
fined to  the  40-125  fathom  zone  between  the 
offings  of  Cape  Lookout,  North  Carolina,  and  of 
Nantucket.  It  seems  to  be  the  most  plentiful  off 
Virginia,  in  the  general  offing  of  Chesapeake  Bay, 
where  considerable  numbers  are  taken  during  the 
winter  trawl  fishing.  They  are  caught  now  and 
then  as  far  as  the  offing  of  Marthas  Vineyard, 
and  Cap'n  Bill  II  trawled  one,  in  July  1952,  south 
of  Nantucket  Lightship,  Lat.  40°02'  N;  Long. 
69°37'  W,  at  75-90  fathoms  which  brings  it  within 
the  arbitrary  boundary  of  the  Gulf  of  Maine. 


Figure  9A. — Chain  dogfish  (.Scyliorhinus  retijer),  male,  about  17  inches  long,  New  Jersey.     After  Bigelow  and  Schroeder. 


SMOOTH  DOGFISHES. 

These  are  rather  small  sharks,  with  two  dorsal 
fins  without  spines,  the  second  dorsal  (in  Atlantic 
species)  nearly  as  large  as  the  first,  and  they  have 
an  anal  fin.  The  tail  fin  is  very  strongly  asym- 
metrical, its  lower  anterior  corner  forming  a  low 
but  rather  definite  lobe  in  some,  but  not  in  others. 
The  teeth  are  small,  with  several  rows  in  function 
imultaneously,  flat,  and  pavement-like  in  some, 


FAMILY  TRIAKIDAE 

but  with  three  or  four  definite  cusps  in  others. 
The  eye  has  no  nictitating  ("winking")  mem- 
brane, but  only  a  longitudinal  fold  along  the 
lower  eyelid.  They  resemble  the  requiem  sharks 
(Family  Carcharhinidae,  p.  36),  except  for  the 
teeth,  and  for  the  lack  of  a  nictitating  membrane. 
Only  one  species  is  known  from  the  Gulf  of  Maine, 
or  is  ever  likely  to  be  found  there. 


Smooth  dogfish  Mustelus  canis  (Mitchill)   1815 

Smooth  dog;  Smooth  hound;  Grayfish 

Bigelow  and  Schroeder,  1948,  p.  244. 

Garman,  1913,  pi.  4,  figs.  6-9,  as  Galeorhinus  laevis. 


FISHES   OF  THE  GULF  OF   MAINE 


35 


Figure  10. — Smooth  dogfish  (Mustelus  canis),  male,  about  31  inches  long,  Woods  Hole.  A,  tooth  band  of  right-hand 
side  of  upper  jaw,  about  1.8  times  natural  size;  B,  teeth  of  another  specimen,  about  6  times  natural  size.  From 
Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


Description. — The  smooth  dog  is  easily  identi- 
fied among  Gulf  of  Maine  sharks  by  having  two 
large  spineless  dorsal  fins,  the  second  only  a  little 
smaller  than  the  first,  combined  with  low,  flat, 
pavement-like  teeth.  So  different,  indeed,  are  its 
teeth  from  the  awl-like  or  blade-like  teeth  of  all 
our  other  sharks  that  a  glance  at  the  mouth  is 
enough  to  separate  this  species  from  the  young  of 
any  larger  Gulf  of  Maine  shark.  In  form  this 
little  shark  is  slender,  flattened  below,  with  taper- 
ing but  blunt  snout.  Its  first  dorsal  fin  originates 
nearly  over  the  hind  angle  of  the  pectorals.  The 
second  dorsal  fin  is  about  twice  as  large  as  the 
anal,  over  which  its  stands.  The  tail  is  of  typical 
"shark"  shape,  i.  e.  with  upper  lobe  much  longer 
than  lower.  The  hind  margin  of  the  upper  lobe 
of  the  caudal  is  deeply  notched  near  the  tip;  the 
lower  caudal  lobe  is  very  small. 

Color. — Upper  surface  grayish  olive,  slaty  gray 
or  brown,  lower  surface  yellowish  or  grayish 
white.  Newborn  specimens  have  the  upper  part 
of  the  first  dorsal  fin  edged  with  dusky  gray;  the 
apex  of  the  second  dorsal  sooty  edged  or  tipped, 
but  with  the  rear  edge  white;  the  tail  fin  with  a 
sooty  blotch  above  near  the  tip,  but  white  edged 
below.  But  these  markings  have  mostly  faded 
out  by  the  time  the  little  "dog"  has  grown  to  a 
length  of  two  feet  or  so.  Smooth  dogs  have  a 
greater  ability  than  most  sharks  to  change  shade 
to  suit  their  surroundings,  paling  to  a  translucent 


pearly  tint  above  white  sand,  but  darkening  on 
dark  bottom.80 

Size. — Smooth  dogs  range  from  about  11% 
inches  to  about  14%  inches  long  when  born.  They 
mature  sexually  at  about  3  feet,  most  of  the  ma- 
ture females  with  young  are  between  about  3 
feet  3  inches  and  4  feet  4  inches  long;  and  a  few 
grow  to  a  length  of  about  5  feet. 

Habits. — The  smooth  dog  is  most  familiar  as  a 
shore  fish  and  a  bottom  swimmer,  commonly 
entering  shoal  harbors  and  bays,  and  even  coming 
into  fresh  water.  But  fishermen  also  report  them 
as  far  offshore  as  the  "tile  fish"  grounds  off 
southern  New  England  and  down  to  a  depth  of 
80  to  90  fathoms.  They  reach  the  northern  part 
of  their  range  only  as  warm-season  visitors;  at 
Woods  Hole  they  arrive  sometime  in  May,  to 
withdraw  in  late  October  or  in  November. 

Food  of  the  smooth  dogfish  consists  chiefly  of 
the  larger  Crustacea,  and  it  is  perhaps  the  most 
relentless  enemy  of  the  lobster,  which  had  been 
eaten  by  no  less  than  16  percent  of  the  fish 
examined  by  Field.  Large  crabs  are  likewise  an 
important  article  in  its  diet,  as  are  the  smaller 
fishes.  It  has  been  estimated  that  10,000  smooth 
dogfish,  in  Buzzards  Bay,  might  devour  more 
than  60,000  lobsters  yearly,  and  perhaps  one-fifth 

*>  Eipcriments  have  shown  that  It  requires  only  1  to  2  hours  for  one  to 
darken,  but  as  much  as  2  days  to  pale  to  the  extreme;  see  Parker  (Biol.  Bull., 
vol.66, 1934,  p.  31). 


36 


FISHERY  BULLETIN   OP   THE    FISH   AND   WILDLIFE   SERVICE 


million  crabs,  besides  a  great  number  of  small 
fish  (menhaden  and  tautog  are  the  species  most 
often  found  in  smooth  dogfish  stomachs).  And 
these  figures  are  based  on  a  sufficient  number  of 
observations  of  the  stomach  contents  to  serve  as  a 
general  indication  of  the  destructiveness  of  the 
smooth  dogfish.  They  also  feed  on  squid,  espe- 
cially in  spring,  and  while  they  do  not  regularly 
take  hard-shelled  mollusks,  razor  clams  have 
been  found  in  the  stomachs  of  several  at  Woods 
Hole.  When  kept  in  captivity  they  are  constantly 
on  the  move,  searching  the  bottom  for  food,  which 
they  find  chiefly  by  the  sense  of  smell  though 
their  sight  is  also  keen.81  Any  crab  that  may  be 
offered  is  soon  found,  seized,  shaken  to  and  fro, 
and  eaten.  And  with  packs  of  these  sea  hounds 
hunting  over  every  square  foot  of  our  southern 
bays  and  sounds  it  is  a  wonder  any  of  the  larger 
crustaceans  escape  where  dogfish  are  abundant. 
Field 82  also  made  the  interesting  observation 
that  the  smooth  dogfish  never  molested  healthy 
and  active  menhaden  but  soon  devoured  any  sick 
or  injured  fish  that  might  be  in  the  same  tank 
with  them. 

As  fhis  is  not  a  characteristic  Gulf  of  Maine 
fish,  we  need  merely  note  that  it  is  one  of  the 
sharks  that  develop  a  placental  connection  be- 
tween the  embryos  and  the  mother.  In  other 
words,  it  is  truly  viviparous.  The  period  of 
gestation  appears  to  be  about  10  months;  off 
southern  New  England  the  young  are  born  be- 
tween early  May  and  mid  July.  The  number  in  a 
litter  usually  is  between  10  and  20,  but  as  few  as 
4  have  been  reported.  A  description  of  the  un- 
born young  is  given  by  Fowler.83 


General  range. — Coastal  waters  of  the  western 
Atlantic,  from  Uruguay  and  southern  Brazil, 
regularly  to  Cape  Cod,  and  to  Passamaquoddy 
Bay  as  a  stray;  also  Bermuda.84 

Occurrence  in  the  Gulf  of  Maine. — The  smooth 
dog  is  the  second  most  numerous  shark  along  the 
southern  coast  of  New  England,  though  falling 
far  short  of  the  spiny  dogfish  (p.  50).  At  Woods 
Hole,  for  example,  pound-net  catches  varied 
during  the  summer  of  1903  from  1  to  41,  averaging 
about  7,  and  catches  up  to  100  have  been  reported 
from  the  vicinity  at  one  time.  Similarly,  catches 
of  5  or  6  on  a  hand  line  are  common  in  a  few  hours' 
fishing,  with  as  many  as  10  to  20  reported.  But 
the  elbow  of  Cape  Cod  and  the  region  of  Nan- 
tucket Shoals  mark  so  definite  a  boundary  to  their 
dispersal  eastward  that  while  they  have  been 
reported  from  Provincetown,  from  various  locali- 
ties within  Massachusetts  Bay,  and  even  from  as 
far  north  as  St.  Andrews  in  the  Bay  of  Fundy, 
where  one  was  caught  in  July  1913,  neither  of  us 
had  ever  seen  one  north  of  Cape  Cod  until  Sep- 
tember 21,  1951,  when  an  angler  (Ellery  Sidney) 
showed  us  a  female  about  3  feet  long  that  he  had 
caught  at  Cohasset,  while  casting  with  an  eel 
skin,  for  striped  bass.  So  far  as  known  its 
occasional  incursions  into  the  Gulf  are  sporadic, 
at  least  they  have  not  been  correlated  with 
unusually  warm  summers  or  with  the  presence  of 
other  southern  fishes.  Neither  has  it  been  re- 
ported by  fishermen  from  Georges  or  Browns 
Banks,  nor  was  it  detected  there  by  the  repre- 
sentatives of  the  Bureau  of  Fisheries  during  the 
trawling  investigations  of  the  years  1912  and  1913 
(p.  60),  or  subsequently. 


REQUIEM  SHARKS.     FAMILY  CARCHARHINIDAE 


This  family,  which  includes  a  large  number  of 
species  in  tropical  and  temperate  seas,  is  charac- 
terized by  a  head  of  normal  shape,  eye  with  a 
nictitating  (winking)  membrane,  tail  with  the 
upper  lobe  considerably  larger  than  the  lower  but 
not  very  long,  2  spineless  dorsal  fins,  the  first 
usually  much  larger  than  the  second  in  most  of 

"  The  senses  of  this  shark  have  been  studied  by  Parker  (Bull.,  U.  S.  Bur. 
of  Fish.,  vol.  29,  1M1,  pp.  43-57),  and  by  Sheldon  (Jour.  Compar.  Neurol, 
and  Psychol.,  vol.  19,  1909,  No.  3,  p.  273). 

,!  Rept.  U.  S.  Coram.  Fish.,  (1906),  1907,  Spec.  Pap.  6,  pp.  14-16. 

"  Occas.  Pap.  Mus.  Zoo].,  I'niv.  Mich.,  No.  56, 1918,  p.  15. 


the  Atlantic  species,85  an  anal  fin,  a  caudal  peduncle 
lacking  lateral  keels,  and  sharp,  bladelike  teeth 
with   a   single   cusp.     All    bear  "living"   young; 


B*  Present  indications  are  that  several  more  or  less  isolated  populations  of 
this  shark  exist,  with  their  areas  of  regular  occurrence  separated  by  wide 
gaps,  where  there  Is  little  or  no  intermingling.  One  of  the  best  known  is 
along  the  Atlantic  coast,  Cape  Cod  to  North  Carolina;  another  centers  in 
the  Gulf  of  Mexico-Caribbean  region;  a  third  is  along  southern  Brazil  and 
Uruguay.  For  further  details,  see  Bigelow  and  Schroeder,  Fishes  Western 
North  Atlantic,  Part  1, 1948,  pp.  250-251. 

w  The  lemon  shark  (Negaprion  brePtrowrU)  of  warmer  waters,  which  has 
been  known  to  stray  to  New  Jersey,  is  an  exception  in  this  respect;  its  second 
dorsal  is  nearly  as  large  as  its  first  dorsal. 


FISHES   OF  THE  GULF  OF   MAINE 


37 


some  have  a  placental  connection  between  mother 
and  embryo,  but  others  do  not. 

Tiger   shark    Galeocerdo    cuvier    (LeSueur)    1822 

Leopard  shark 

Bigelow  and  Schroeder,  1948,  p.  266. 

Description. — The  tiger  shark  is  characterized 
among  the  Atlantic  members  of  its  family  by  the 
forward  position  of  its  first  dorsal  fin  (origin  about 
over  the  arm  pit  of  the  pectorals),  combined  with 
a  caudal  peduncle  with  a  low  longitudinal  ridge 
of  skin  on  either  side,  besides  a  well-marked 
semilunar  pit  below  as  well  as  above;  a  very  small 
second  dorsal  fin;  a  furrow,  about  as  long  as  the 
snout  along  either  side  of  the  upper  jaw;  a  very 
slender-tipped  caudal  fin  with  moderately  large 
and  pointed  lower  lobe;  and  large  teeth  alike  in 
the  two  jaws,  of  very  characteristic  shape,  with 
convex  inner  margins,  deeply  and  conspicuously 
notched  outer  margins  and  strongly  serrate  edges 

(%.  ID- 

Young  tiger  sharks  are  rather  slender,  but 
they  become  very  heavy  forward,  with  growth, 
though  they  continue  tapering  toward  the  tail. 
The  first  dorsal  fin  is  high,  triangular,  and  nearly 
as  large  as  the  pectorals,  while  the  second  dorsal 
is  hardly  one-third  to  one-fourth  as  high  as  the 
first  and  stands  over  the  anal,  which  is  of  about 
equal  size.  The  lower  tail  lobe  is  almost  half  as 
long  as  the  upper,  the  rear  margin  of  which  is 
notched  near  the  tip.  The  large  size  of  the 
head,  with  very  short,  obtusely  rounded  front 


outline,  and  broad  mouth  occupying  nearly 
four-fifths  of  the  width  of  the  head,  with  long 
grooves  along  the  upper  jaw,  combined  with  the 
unique  shape  of  its  teeth,  make  the  "tiger"  easy 
to  recognize  among  Gulf  of  Maine  sharks. 

Color. — Gray,  or  grayish  brown,  darkest  on 
the  upper  surface.  Young  "tigers"  up  to  5  or  6 
feet  long,  are  more  or  less  conspicuously  spotted 
or  barred  with  darker  brown  on  the  back  and  along 
the  upper  parts  of  the  sides.  But  these  markings 
fade  with  advancing  age  until  large  specimens 
are  plain  colored,  or  nearly  so. 

Size. — Tiger  sharks  are  small  at  birth,  corre- 
sponding to  the  large  numbers  in  a  litter,  free 
living  specimens  having  been  reported  only 
18  to  19  inches  long.  By  the  time  they  mature 
they  are  among  the  larger  sharks;  but  their  size 
has  often  been  overestimated.  The  majority 
of  tigers  caught  in  centers  of  abundance  are  less 
than  12  to  13  feet  long,  and  the  largest  measured 
lately  in  the  western  Atlantic  was  one  of  about 
18  feet,  from  Cuba.  Repeated  statements  that 
the  tiger  grows  to  a  maximum  length  of  30  feet 
have  no  reliable  foundation,  so  far  as  we  can 
discover. 

A  4-foot  specimen  from  Woods  Hole  weighed 
25%  pounds  when  taken  from  the  water.  Larger 
tigers  vary  widely  in  weight  at  given  lengths 
depending  on  how  fat  they  are  and  on  the  stage 
of  development  of  the  young  in  gravid  females. 
Specimens  from  various  localities  have  weighed 
37  pounds  at  5%  feet;  168  pounds  at  6  feet;  366  to 
718  pounds  at  10  to  11  feet;  450  to  825  pounds 
at  11  to  12  feet;  630  to  1,324  pounds  at  12  to  13 


Figure  11. — Tiger  shark  (Galeocerdo  cuvier),  young  male,  about  49  inches  long,  Rhode  Island.     A,  upper  tooth,  and 
B,  lower  tooth  of  larger  specimen,  enlarged.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


38 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


feet;  and  1,028  to  1,395  pounds  at  13  to  14  feet.88 
Habits. — This  voracious  shark,  with  wide  jaws 
and  powerful  teeth,  preys  upon  the  large  sea 
turtles,  other  sharks,  fish,  and  occasionally  on 
invertebrates  such  as  horseshoe  crabs,  crabs, 
conchs,  whelks.  It  is  proverbial  for  its  habit 
of  feeding  on  slaughter-house  wastes  or  any  other 
carrion.  Remnants  of  squeteague,  mackerel, 
hake,  scup,  menhaden,  goosefish,  and  dogfish  all 
have  been  found  in  stomachs  of  tiger  sharks  taken 
at  Woods  Hole.87  There  is  no  placental  connec- 
tion between  mother  and  young,  and  the  broods 
are  very  large,  as  many  as  82  having  been  counted 
in  a  large  female;  but  other  litters  as  small  as 
10  to  14.  In  the  West  Indies  it  is  much  dreaded, 
whether  or  not  with  good  cause. 

General  range. — Cosmopolitan  in  the  warmer 
waters  of  all  oceans;  straying  northward  as  far 
as  Cape  Cod  on  the  American  coast  of  the  Atlantic. 
Occurrence  in  the  Gulf  of  Maine. — A  few  tiger 
sharks  are  taken  in  fish  traps  in  the  Woods  Hole 
region  every  year,  seldom  before  August  or  later 
than  October  although  one  was  caught  there  July  20, 
1951.88  These  specimens  usually  have  been  about 
5  feet  long,  at  most  about  8  feet,  and  very  rarely 
does  a  full-grown  tiger  shark  stray  so  far  from 
its  tropical  home.  The  tiger  has  not  yet  been 
recorded  (on  reliable  evidence)  from  within  the 
limits  of  the  Gulf  of  Maine.     It  is  included  here 

M  For  further  details  and  references,  see  Bigelow  and  Schroeder,  Fishes 
Western  North  Atlantic,  Pt.  1, 1948,  p.  269. 

»'  Bell  and  Nichols  (Copela,  No.  92,  March  1921,  pp.  17-20)  list  the  stomach 
contents  of  a  number  of  tiger  sharks  caught  off  Morehead  City,  N.  C. 

"  This  shark  was  8  feet,  3  Inches  long,  taken  In  a  pound  net  off  Quisset 
Harbor,  Buzzards  Bay. 


because  of  the  likelihood  that  a  stray  specimen 
may  occasionally  round  the  elbow  of  Cape  Cod, 
or  be  encountered  on  the  offshore  Banks.89 

Blue  shark  Prionace  glauca  (Linnaeus)  1758 

Blue  dog 

Bigelow  and  Schroeder,  1948,  p.  282. 

Garman,  1913,  pi.  3,  figs.  1-3  (as  Galeus  glaucus). 

Description. — The  blue  shark  is  slender-bodied, 
thickest  about  its  mid-length,  and  tapers  toward 
head  and  tail  (a  shape  usually  named  "fusiform"). 
Its  snout  is  long  with  rounded  tip.  Its  first  dorsal 
fin  is  of  moderate  size,  standing  far  back  with  the 
mid  point  of  its  base  about  midway  between  the 
inner  corners  of  the  pectorals  (when  these  are  laid 
back)  and  the  points  of  origin  of  the  pelvic  fins. 
The  second  dorsal  fin  is  less  than  one-half  as  high 
as  the  first,  and  is  about  equal  in  size  to  the  anal 
over  which  it  stands.  The  pectorals  are  narrow 
and  very  long,  their  tips  reaching  back  nearly  as 
far  as  the  rear  corner  of  the  first  dorsal.  The 
lower  lobe  of  the  caudal  fin  (measured  along  its 
anterior  edge)  is  about  one-half  as  long  as  the 
upper  lobe;  the  latter  is  conspicuously  notched 
near  the  tip,  and  both  of  the  lobes  of  the  caudal 
fin  are  slender  tipped. 

The  teeth  are  large,  sharp-pointed,  with  serrate 
edges,  and  distinctive  in  shape.  The  uppers  are 
so  closely  spaced  that  the  bases  of  adjacent  teeth 

"  The  statement  In  the  first  edition  that  a  tiger  shark  was  once  taken  at 
Provlncetown  was  an  error.  The  original  description  of  the  specimen  In 
question  (Atwood,  Proc.  Boston  Soc.  Nat.  Hist.,  vol.  10, 1865,  p.  81)  suggests 
that  it  was  a  mako  (Isurus  oxyrinchus) . 


Figure  12. — Blue  shark  (Prionace  glauca),  male,  about  7  feet  2  inches  long,  off  Marthas  Vineyard.  A,  third  left-hand 
upper  tooth,  counted  from  mid-point  of  jaw;  B,  ninth  left-hand  upper  tooth;  C,  third  left-hand  lower  tooth;  and 
D,  eighth  left-hand  lower  tooth;  about  1.6  times  natural  size.  From  Bigelow  and  Schroeder.  Drawings  by  E.  N. 
Fischer. 


FISHES   OP  THE  GULF   OF   MAINE 


39 


overlap.  The  median  upper  tooth  is  nearly 
symmetrical,  but  those  along  the  sides  of  the 
mouth  have  strongly  convex  outer  margins,  and 
deeply  concave  inner  margins,  while  their  points 
curve  sharply  outward  toward  the  respective 
corner  of  the  mouth.  The  lower  teeth  are  nar- 
rower, more  nearly  symmetrical,  and  nearly 
erect. 

Color. — Living  specimens  are  dark  indigo  blue 
along  the  back,  shading  to  a  clear  bright  blue  90 
along  the  sides;  but  this  beautiful  hue  changes  to 
a  slaty  or  sooty  gray  soon  after  death.  The  lower 
surface  is  snow-white,  but  with  the  tips  of  the 
pectorals  dusky  and  the  anal  fin  partly  sooty. 

Size. — The  usual  length  at  birth  seems  to  be 
between  Y%  and  2  feet.81  Blue  sharks  do  not  ma- 
ture until  they  have  grown  to  be  7  or  8  feet  long, 
to  judge  from  the  sizes  of  the  females  that  have 
been  found  with  young;  the  longest  we  have 
handled  was  almost  exactly  11  feet  long.  The 
fact  that  the  greatest  measured  length  so  far  re- 
liably reported  was  only  12  feet  7  inches  (3.83 
meters)  suggests  that  repeated  characterizations 
of  the  blue  shark  as  commonly  growing  to  15  feet 
are  an  exaggeration.  If  any  grow  to  20  feet,  as 
is  rumored,  they  must  be  giants  of  their  kind. 

Remarks. — The  very  long  slender  pectorals  of 
the  blue  shark,  combined  with  its  long  narrow 
snout,  the  position  of  its  first  dorsal  fin  far  back, 
and  its  brilliant  blue  color,  give  it  an  aspect 
very  different  from  that  of  the  tiger  shark  (p.  37), 
of  the  sharp-nosed  shark  (p.  40),  the  dusky  or 
brown  sharks  (pp.  41-43),  or  that  of  any  other 
carcharhinid  shark  that  might  perhaps  straggle  to 
the  Gulf  of  Maine. 

Habits. — The  blue  shark  is  "encountered  indif- 
ferently far  out  at  sea  and  in  continental  waters, 
its  wanderings  no  doubt  directed  chiefly  by  the 
search  for  food,  though  it  may  drift  with  ocean 
currents.  It  is  frequently  seen  at  the  surface, 
swimming  lazily  with  first  dorsal  fin  and  tip  of 
caudal  out  of  water,  or  basking  in  the  sun.  There 
is  no  reason  to  suppose  that  it  ever  descends  to 
any  great  depth."  92  They  sometimes  follow  sail- 
ing ships  for  days  on  end,  to  pick  up  scraps,  and 
their  habit  of  gathering  when  a  sperm  whale  was 


killed,  to  feed  on  the  carcass,  was  proverbial  dur- 
ing the  days  of  the  sperm  whale  fishery.93  But 
their  normal  diet  is  smaller  fishes,  of  whatever 
kinds  may  be  available.  In  northern  waters 
herring,  mackerel,  spiny  dogfish,  and  various 
others  have  been  found  in  their  stomachs.  And 
we  have  several  times  seen  a  blue  shark  pick  up 
a  tagged  cod,  haddock  or  American  pollock  that 
we  had  put  back  in  the  water,  on  Georges  Bank. 

The  blue  shark  is  viviparous,  that  is  to  say,  the 
embryo  has  a  well  developed  placenta  attached  to 
the  mother.  As  many  as  28  to  54  young  have 
been  reported  in  a  litter  in  the  Mediterranean. 

General  range. — Cosmopolitan  on  the  high  seas 
in  the  warmer  parts  of  all  the  oceans,  including 
the  Mediterranean;  ranging  northward  to  outer 
Nova  Scotia  and  as  a  stray  to  the  Banks  of  New- 
foundland in  the  western  side  of  the  Atlantic;  to 
England  and  Scotland  in  the  east,  with  stray 
specimens  reaching  the  Orkneys  and  southern 
Norway.  This,  we  think,  is  by  far  the  most  nu- 
merous of  the  large,  oceanic  sharks;  it  is  the  one 
with  which  the  sperm  whalers  were  the  most 
familiar;  the  one  around  which  many  of  the  super- 
stitions about  sharks  have  developed;  and  the  one 
with  which  we  have  had  to  do  most  often. 

Occurrence  in  the  Gulf  oj  Maine  and  along  Nova 
Scotia. — Only  one  blue  shark  had  been  reported 
definitely  from  the  Gulf  of  Maine  in  scientific  lit- 
erature, up  to  the  time  the  first  edition  of  this  book 
was  printed,  though  it  was  known  to  be  rather  com- 
mon along  outer  Nova  Scotia.  But  we  have 
learned  since  then  that  it  is  a  regular  summer  visi- 
tor to  the  southern  and  western  parts  of  the  Gulf, 
appearing  occasionally  in  July,  more  often  in  Aug- 
ust and  September.  In  1928,  for  example,  we 
caught  one  on  Stellwagen  Bank  on  August  26,  saw 
one  over  the  northern  end  of  Jeffreys  Ledge  on 
September  2,  and  caught  four  on  Platts  Bank  on 
September  3,  with  others  in  sight  from  the  vessel  at 
nearly  all  times  throughout  the  day.  And  many 
more  have  been  seen  or  caught  subsequently,  on 
Platts  Bank,  in  Massachusetts  and  Cape  Cod  Bays, 
where  18  were  reported  to  us  during  the  summer  of 
1935,94  on  Georges  Bank  where  blue  sharks,  swim- 
ming at  the  surface,  are  a  familiar  sight  in  summer; 
and  on  Browns  Bank.     Two  have  also  been  re- 


"  "Sailor  blue,"  as  shown  In  Rldgeway's  Color  Standards  and  Color 
Nomenclature,  1912,  p.  21. 

81  Embryos  have  been  reported  as  long  as  about  17H  inches,  and  free-living 
specimens  as  small  as  20-21  inches. 

"Blgelow  and  Schroeder,  Fishes  Western  North  Atlantic,  Pt.  1,  1948,  p. 
286. 


»  Nichols  and  Murphy  (Brooklyn  Mus.  Sci.  Bull.,  vol.  3,  No.  1, 1916,  p. 
9)  have  given  a  graphic  account  of  the  blue  shark  as  it  was  met  with  by 
whalers  on  the  high  seas. 

"  By  J.  R.  Lowes,  an  experienced  shark  fisherman. 


40 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


ported  to  us  recently  from  the  coast  of  Maine,  a  few 
miles  east  of  Casco  Bay.95 

We  have  never  heard  of  a  blue  shark  in  the  north- 
eastern corner  of  the  Gulf,  in  the  Bay  of  Fundy, 
nor  along  western  Nova  Scotia,  whence  they  may 
be  barred  by  colder  surface  waters.  But  fisher- 
men are  familiar  with  tbem  off  the  outer  coast  of 
Nova  Scotia,  both  offshore,  and  also  near  the  coast 
at  the  times  when  the  warm  surface  water  presses 
inshore. 

Blues  were  reported  near  Halifax,  for  instance, 
from  time  to  time  between  August  15  and  October 
10,  1920,  some  coming  close  in  to  the  entrance  to 
the  Harbor.  And  two  specimens  have  been  re- 
ported at  Canso,96  but  whether  the  "blue  dogs" 
described  by  local  fishermen  as  common  on  the 
neighboring  banks  actually  are  this  shark,  or  per- 
haps the  porbeagle,  seems  doubtful.  It  has  also 
been  recorded  from  the  southwest  part  of  the 
Grand  Bank  of  Newfoundland.97 

Following  westward  from  Cape  Cod,  we  find 
many  records  of  blues  from  the  traps  near  Woods 
Hole,  and  they  are  often  seen  (or  harpooned)  on 
the  continental  shelf  in  the  offing.  Twenty-eight 
were  counted  4  to  10  miles  off  Block  Island  for  ex- 
ample, during  one  hour,  and  something  like  150  to 
200  during  the  day  (13  of  them  were  harpooned)  on 
August  22,  1943. 

Most  of  the  blues  that  are  seen  or  taken  off  our 
northern  coast  are  medium  sized  or  larger,  though 
very  small  ones  are  taken  from  time  to  time.98 

"  By  the  late  Walter  H.  Rich,  who  was  long  associated  with  the  U.  S.  Bu- 
reau of  Fisheries. 

«  Cornish,  Contr.  Canadian  Biol.  (1902-1905)  1907,  p.  81. 

«  Rept.  Newfoundland  Fish.  Res.  Lab.,  1935,  p.  79. 

88  Robert  Goffin  reports  one  only  20  Inches  long,  from  Menemsha  Bight, 
near  Woods  Hole,  August  31 ,  1925;  we  have  seen  one  of  21  Inches,  taken  a  few 
miles  off  Block  Island,  August  22, 1943;  and  F.  D.  Firth  reports  one  34H  Inches 
long  taken  65  miles  southeast  of  Highland  Light,  Cape  Cod,  on  October  23, 
1930. 


And  for  some  obscure  reason  all  but  two  of  the 
adults  seen  in  our  Gulf,  for  which  we  have  the  per- 
tinent information,  have  been  males. 

Commercial  importance. — This  shark  is  of  no 
commercial  value.  A  few  are  caught  by  anglers, 
mostly  on  natural  bait,  and  a  Blue  will  sometimes 
take  an  artificial  lure;  we  hooked  one  off  Boone 
Island,  Maine,  on  a  feather  jig,  tipped  with  pork 
rind.  We  have  never  bad  blues  put  up  much  re- 
sistance on  a  heavy  hand  line  until  hauled  in  to  the 
side  of  the  vessel,  when  they  thrash  about  vio- 
lently, but  it  is  said  that  a  large  one  will  make  long 
and  powerful  runs,  if  hooked  on  rod  and  reel. 

The  blue  shark  has  always  been  looked  on  with 
contempt  by  the  sperm  whalers,  who  were  more 
familiar  with  it  than  anyone  else.  We  find  no  well- 
authenticated  case  of  one  attacking  a  swimmer, 
sailors'  yarns  to  the  contrary  notwithstanding. 

Sharp  nosed  shark  Scoliodon  terrae-novas 
(Richardson)  1836 

Bigelow  and  Schroeder,  1948,  p.  295. 
Garman,  1913,  pi.  2,  figs.  1-4. 

Description. — This  little  shark  is  separable  from 
any  other  carcharhinid  that  has  yet  been  reported 
from  the  Gulf  of  Maine  or  that  is  likely  to  be,  by  its 
upper  and  lower  teeth  which  are  perfectly  smooth 
along  the  edges  from  tip  to  base,  combined  with  a 
so-called  "labial  furrow"  of  considerable  length 
running  forward  along  each  side  of  each  jaw  from 
the  corner  of  the  mouth  toward  the  nostril. 
This  last  character,  while  not  conspicuous,  is  a 
precise  one. 

The  trunk  is  slender,  highest  about  at  the  first 
dorsal  fin,  tapering  both  fore  and  aft.  The  snout 
varies  rather  widely  in  length  and  in  bluntness  at 
the  tip.     The  point  of  origin  of  the  first  dorsal  fin 


Figure  13. — Sharp-nosed  shark  (Scoliodon  lerrae-novae) ,  female,  about  31  inches  long,  from  the  Bahamas.     From  Bigelow 

and  Schroeder.     Drawing  by  E.  N.  Fischer. 


FISHES   OF  THE  GULF  OF   MAINE 


41 


is  about  over  the  inner  corners  of  the  pectorals 
when  the  latter  are  laid  back;  its  height  is  about 
one-half  as  great  as  the  distance  from  the  tip  of 
the  snout  to  the  level  of  the  origin  of  the  pectorals. 
The  second  dorsal  is  only  about  one-quarter  as 
high  as  the  first;  its  point  of  origin  is  about  over 
the  mid-point  of  the  base  of  the  anal  fin ;  the  anal 
is  a  little  larger  than  the  second  dorsal.  The  tail 
fin  occupies  about  one-quarter  of  the  total  length 
of  the  shark;  its  lower  lobe  (measured  along  the 
anterior  edge)  is  a  little  less  than  one-half  as  long 
as  the  upper  lobe,  the  rear  edge  of  which  is  deeply 
notched  near  the  tip.  The  pectoral  fins  are  smaller 
relatively  than  in  any  other  local  species  of  this 
family,  their  length,  armpit  to  tip,  being  only  a 
little  greater  than  the  height  of  the  first  dorsal 
fin.  The  teeth  are  alike  in  shape  in  the  two  jaws, 
sharp-pointed  and  smooth  edged;  those  in  the 
center  of  the  mouth  are  symmetrical  and  erect, 
but  those  along  the  sides  have  weakly  concave 
inner  margins,  but  deeply  notched  outer  margins, 
and  are  increasingly  oblique  toward  the  corners 
of  the  mouth. 

Color. — Brown  to  olive  gray  above,  with  the  dorsal 
and  caudal  fins  more  or  less  dark  edged;  white 
below  and  along  the  rear  margins  of  the  pectorals. 

Size. — Mature  specimens  are  commonly  between 
26  and  30  inches  long;  a  few  grow  to  36  inches. 

General  range. — Both  sides  of  the  tropical-sub- 
tropical Atlantic;  Morocco  to  Cameroon  and  the 
Cape  Verde  Islands  in  the  east ;  Uruguay  to  North 
Carolina  in  the  west;  occasional  to  Woods  Hole, 
and  as  a  stray  to  the  Bay  of  Fundy. 

Occurrence  in  the  Gulf  of  Maine. — Our  only 
reason  for  including  this  warm-water  shark  is 
that  one  was  taken  at  Grand  Manan  Island,89  at 
the  mouth  of  the  Bay  of  Fundy,  in  1857.1""2 

Early  reports  of  it  from  Newfoundland  were 
based  on  a  misconception. 

Sharks  of  the  Genus  Carcharhinus 

The  members  of  the  genus  Carcharhinus  are  set 
apart  from  other  Atlantic  members  of  the  family 
Carcharhinidae  by  the  following  combination  of 
characters :  The  mid-point  of  base  of  the  first  dorsal 
fin  is  at  least  as  near  to  the  level  of  the  axils  of 
the  pectorals  as  to  the  level  of  the  origin  of  the 

"  This  specimen,  collected  by  A.  E.  Verrill,  Is  in  the  Museum  of  Com- 
parative Zoology. 

1-1  See  Jordan  and  Evermarm,  Bull.  47,  V.  S.  Nat.  Mus.,  Pt.  1,  1896,  p.  43, 
footnote. 


pelvics  (separating  them  from  the  blue  shark, 
p.  38) ;  no  labial  furrows  on  lower  jaw,  and  furrow 
on  upper  jaw  reduced  to  a  very  short  slit  at  the 
extreme  corner  of  the  mouth,  directed  outward 
(separating  them  from  the  tiger  shark,  p.  37,  and 
from  the  sharp-nosed  shark,  p.  40) ;  second  dorsal 
fin  much  smaller  than  first  dorsal  (separating 
them  from  the  lemon  shark,  p.  35,  footnote  85); 
edges  of  upper  teeth  more  or  less  finely  serrate  but 
without  larger  denticles  near  the  base,  and  edges 
of  lower  teeth  perfectly  smooth,  without  lateral 
denticles  (separating  them  from  the  tiger  shark, 
p.  37,  from  the  sharp-nosed  shark,  p.  40),  and 
from  Paragaleus  pecloralis,  a  tropical  shark  that 
has  been  taken  off  southern  New  England.3 

This  is  a  warm-water  group,  fifteen  species  of 
which  are  known  to  inhabit  the  western  side  of 
the  Atlantic,  most  of  them  resembling  one  another 
closely  in  general  aspect.  Only  one  of  these  (the 
dusky  shark,  described  on  p.  41)  has  yet  been 
reported  reliably  from  within  the  confines  of  the 
Gulf  of  Maine,  while  only  one  other  (the  brown 
shark,  p.  43)  is  likely  to  be  found  there.  If  a 
stray  Carcharhinus  from  offshore  that  does  not 
agree  with  the  following  descriptions  of  one  or 
other  of  these  should  be  taken  on  Georges  Bank,  or 
on  Nantucket  Shoals  east  of  the  longitude  of 
Cape  Cod,  we  hope  that  its  captor  can  identify 
it  by  means  of  the  keys  and  descriptions  of  the 
genus  that  we  have  given  in  Part  1  of  the  Fishes 
of  the  Western  North  Atlantic. 

Dusky   shark    Carcharhinus  obscurus   (LeSueur) 

1818. 

Bigelow  and  Schroeder,  1948,  p.  382. 

Description. — The  combination  of  characters 
that  place  the  dusky  shark  among  the  western 
Atlantic  members  of  its  genus  are:  Trunk  about 
one-fifth  as  high  at  first  dorsal  fin  as  it  is  long  to 
origin  of  the  caudal  fin,  tapering  both  forward 
and  rearward;  snout  broadly  rounded  in  front,  its 
length  in  front  of  the  nostrils  less  than  the  distance 
between  the  nostrils;  the  front  edge  of  the  nostril 
is  not  expanded  as  a  definite  lobe;  the  midline  of 
the  back  between  the  two  dorsal  fins  has  a  low 
but  definite  ridge,  a  character  which  is  very  pre- 
cise, though  seemingly  minor;  the  first  dorsal  fin  is 
considerably  smaller   than   in   the   brown  shark 

'  For  description,  see  Bigelow  and  Schreeder,  Fishes  of  the  Western  North 
Atlantic,  Pt.  1, 1948,  p.  276. 


42 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  14. — Above:  Dusky  shark  (Carcharhinus  obscurus),  female  about  39  inches  long,  Woods  Hole.  A,  third  upper 
tooth;  B,  fourth  lower  tooth;  C,  ninth  upper  tooth;  D,  tenth  lower  tooth;  about  2.4  times  natural  size.  Below: 
Brown  shark  (Carcharhinus  milberti),  female,  about  4  feet  10  inches  long,  from  Woods  Hole.  A,  ninth  upper  tooth; 
B,  eighth  lower  tooth;  C,  third  lower  tooth;  about  1.4  times  natural  size.  From  Bigelow  and  Schroeder.  Drawings 
by  E.  N.  Fischer. 


(p.  43),  with  more  deeply  concave  rear  margin, 
its  point  of  origin  about  over  the  inner  corner  of 
the  pectoral  (over  the  armpit  of  the  pectoral  in 
the  brown  shark);  its  apex  is  narrowly  rounded. 
The  free  rear  corner  of  the  second  dorsal  fin  is  less 
than  twice  as  long  as  the  vertical  height  of  the 
fin.  The  anal  fin  is  a  little  longer,  along  the  base, 
than  the  second  dorsal  and  stands  about  under  the 
latter.  The  caudal  fin  occupies  between  one* 
quarter  and  one-third  of  the  total  length  of  the 
shark,  the  lower  caudal  lobe  (measured  along  it9 
anterior  edge)  is  about  two-fifths  as  long  as  the 
upper  lobe;  and  the  upper  lobe  is  noticeably 
slender  toward  its  tip.  The  pectorals  are  about 
as  long  (from  origin  to  tip)  as  the  distance  from 


the  tip  of  the  snout  to  the  level  of  the  first  pair  of 
gill  openings,  usually  narrower,  relatively,  than 
in  the  brown  shark,  and  sometimes  more  definitely 
sickle-shaped. 

The  upper  teeth  are  broadly  triangular;  nearly 
erect  toward  the  center  of  the  mouth  but  weakly 
oblique  toward  its  corners;  their  inner  margins 
are  nearly  straight,  the  outer  margins  increasingly 
concave  outward  along  the  jaw.  The  lower 
teeth  are  erect,  symmetrical,  with  narrow  cusp  on 
a  broadly  expanded  base.  Both  the  upper  teeth 
and  the  lower  are  serrate  along  the  edges,  the 
lower  the  more  finely  so. 

Color. — All  the  fresh  caught  specimens  we  have 
seen  have  been  bluish  or  leaden  gray  on  the  back 


FISHES    OF   THE   GULF   OF    MAINE 


43 


and  upper  part  of  the  sides,  including  the  pectorals, 
but  this  shark  has  also  been  described  as  pale 
gray  above  or  even  dirty  white,  perhaps  over  a 
white  sand  bottom.  The  trunk  is  white  below, 
the  pectorals  grayish,  darkening  to  sooty  at  their 
tips;  the  pelvics  and  anal  fins  grayish  white. 

Size. — -The  usual  length  at  birth  is  a  little  more 
than  three  feet.*  Adult  dusky  sharks  so  far 
measured  have  ranged  from  10  feet  4  inches  to 
11  feet  8  inches  in  length,  and  they  are  said  to 
grow  to  14  feet,  though  perhaps  not  on  very 
convincing  evidence. 

General  range. — Western  Atlantic,  north  to 
southern  New  England  and  to  Georges  Bank, 
south  to  southern  Brazil,  at  least  by  name.  A 
shark  very  closely  allied  to  obscurus  has  been 
reported  under  that  name  in  the  eastern  Atlantic, 
from  Spain  to  Table  Bay,  South  Africa,  including 
Madeira,  the  Canaries,  the  Cape  Verdes,  Ascen- 
sion Island,  and  St.  Helena.  But  we  have  yet 
to  learn  its  precise  relationship  to  the  obscurus  of 
the  western  Atlantic. 

Occurrence  in  the  Gulf  of  Maine. — The  dusky 
shark  has  been  taken  repeatedly  off  the  coasts  of 
New  Jersey  and  of  Long  Island,  N.  Y.;  also  at 
Woods  Hole,  where  we  have  handled  12  specimens 
during  the  past  few  summers,  6  of  them  in  August 
1944.  But  it  so  seldom  strays  to  cooler  waters 
farther  east  that  only  one  shark  has  been  recorded 
from  Nantucket,  and  one  from  Georges  Bank, 
that  probably  were  of  this  species  and  not  some 
other  carcharhinid.6  Thus  it  has  no  real  place  in 
the  fauna  of  the  Gulf.6 

Brown  shark  Carcharhinus  milberti  (Miiller  and 
Henle)  1841 

Sand  bar  shark 

Bigelow  and  Schroeder,  1948,  p.  368. 
Garman,  1913,  pi.  3,  figs.  4-6  (as   Carcharinus  platyo- 
don). 

Description. — The  brown  shark  differs  from  the 
dusky  (only  member  of  its  genus  that  seems 
actually  to  have  been  taken  within  the  Gulf)  in 


the  more  forward  position  and  larger  size  of  its 
first  dorsal  fin,  in  its  broader  pectorals,  and  in  its 
stouter  trunk,  heaviest  forward  (compare  speci- 
mens in  figure  14).  Also,  the  anterior  edge  of  its 
nostril  is  expanded  as  a  low  but  definite  triangular 
lobe,  which  is  not  the  case  in  the  dusky  shark. 
Other  characters  (in  combination)  that  mark  it 
off  from  other  members  of  this  genus  that  might 
stray  to  the  Gulf  are:  Mid-line  of  the  back  with 
a  low  ridge  between  the  two  dorsal  fins;  snout 
forward  of  a  line  connecting  the  front  margins  of 
the  nostrils,  considerably  shorter  than  the  distance 
between  the  nostrils;  point  of  origin  of  second 
dorsal  fin  about  over  origin  of  anal  fin,  its  free 
rear  corner  only  a  little  longer  than  the  height 
of  the  fin;  apex  of  first  dorsal  fin  angular;  length 
of  pectorals  along  anterior  margin  about  as  great 
as  distance  from  tip  of  snout  to  level  of  second 
pair  of  gfil  openings;  distance  from  rear  tips  of 
pelvic  fins  to  origin  of  anal  fin  as  long  as  base  of 
anal  fin,  or  longer,  fifth  gill  openings  longer  than 
horizontal  diameter  of  eye. 

The  teeth  resemble  closely  those  of  the  dusky 
shark  (see  figure  14). 

Color. — Upper  surface  slate  gray  to  brown; 
lower  surface  a  paler  tint  of  the  same  hue,  or 
white;  fins  without  any  conspicuous  black  mark- 
ings. When  alive  some  of  the  dermal  denticles 
are  bright  blue,  at  least  on  some  specimens. 

Size. — Sexual  maturity  is  reached  at  a  length 
of  about  6  feet;  maximum  length  about  eight  feet.7 

General  range. — Southern  Brazil,  Louisiana,  both 
coasts  of  Florida,  and  northward  along  the  Atlantic 
coast  of  the  United  States  to  southern  New 
England;  also  the  tropical-subtropical  belt  of  the 
eastern  Atlantic,  and  the  Mediterranean,  or 
represented  there  by  an  extremely  close  relative.8 

Occurrence  in  the  Gulf  oj  Maine. — Next  to  the 
sand  shark,  this  is  the  most  numerous  of  the 
larger  sharks  along  the  coasts  of  New  Jersey  and 
of  New  York.  Some  visit  the  vicinity  of  Woods 
Hole,  though  so  few  that  the  number  taken  there 
in  most  summers  probably  is  not  greater  than  six 
or  seven.     It  has  not  been  reported  as  yet  from 


*  Embryos  have  been  reported  up  to  38  In.  long  (965  mm.),  and  a  free  living 
specimen  of  only  39  In.  (993  mm.) ;  see  Bigelow  and  Schroeder,  Fishes  Western 
North  Atlantic,  Pt.  1, 1948,  p.  387. 

»  Probably  this  species  and  not  the  brown  shark  because  11-12  feet  long. 

I  In  the  first  edition  of  this  book,  the  dusky  shark  was  said  to  have  been 
taken  at  three  localities  within  the  Gulf.  But  one  of  these  records,  at  least, 
was  almost  certainly  based  on  a  blue  shark,  and  the  others  probably  were 
(Bigelow  and  Schroeder,  Fishes  Western  North  Atlantic,  Pt.  1,  1948,  pp. 
292,  368). 


'  Seven  feet  10  Inches  is  the  greatest  measured  length  that  we  have  found 
recorded,  with  convincing  evidence  that  the  specimen  actually  was  one  of 
this  species. 

•If  the  eastern  Atlantic-Mediterranean  form  Is  actually  Identical  with 
the  American,  as  seems  to  be  the  case,  the  specific  name  milberti  of  Miiller 
and  Henle,  1841,  must  be  replaced  by  plumbeus  proposed  by  Nardo  In  1827 
for  the  brown  shark  of  the  Adriatic. 


44 


FISHKRY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


within  the  limits  of  our  Gulf,  but  is  included  here 
on   the   chance   that   a  stray  specimen  may  be 


taken,  either  on  the  outer  coast  of  Cape  Cod, 
on  Nantucket  Shoals,  or  on  Georges  Bank. 


THE  HAMMER-HEADED  SHARKS.    FAMILY  SPHYRNIDAE 


The  peculiar  hammer-shaped  head,  with  eyes 
far  apart,  sufficiently  characterizes  the  Gulf  of 
Maine  sharks  of  this  family,  which  resembles  the 
requiem  sharks  (p.  36)  otherwise.  Five  species 
are  known  in  the  western  Atlantic,  all  of  them 
tropical-subtropical  in  nature.  Two  of  these  have 
been  reported  from  our  Gulf,  but  only  as  strays. 

Shovelhead  Sphyrna  tiburo  (Linnaeus)  1758 

Bonnet  head  shark 

Bigelow  and  Schroeder,  1948,  p.  420. 

Garman,  1913,  pi.  1,  figs.  4-6  (as  Cestracion  tiburo). 

Description. — The  peculiar  shovel-shaped  head 
of  this  shark  is  enough  to  distinguish  it  readdy 
from  any  other  shark  known  from  the  Gulf  of 
Maine,  except  for  the  hammerhead,  from  which  it 
is  readdy  distinguished  by  the  fact  that  its  head 
is  considerably  narrower,  is  more  rounded  in  front, 
and  is  not  deeply  indented  opposite  each  nostril; 
that  the  posterior  margin  of  its  anal  fin  is  only 
weakly  concave,  and  that  the  outermost  four  or 
five  of  its  lower  teeth  next  each  outer  corner  of  its 
mouth  are  low  and  rounded,  not  blade-like.  The 
eyes  of  the  shovel-head  shark,  like  those  of  the 


hammerhead,  stand  at  either  edge  of  the  expanded 
head;  the  first  dorsal  fin  originates  a  little  behind 
the  "armpit"  of  the  pectoral,  is  somewhat  higher 
than  the  pectorals  are  long,  and  is  higher  than 
long;  the  very  small  second  dorsal  fin  originates  a 
little  behind  the  origin  of  the  anal  fin;  the  upper 
lobe  of  the  tad  is  notably  long  (about  one-third  as 
long  as  the  body  of  the  fish)  and  deeply  notched 
near  the  tip,  the  lower  lobe  is  about  one-third  as 
long  as  the  upper  lobe.  The  anal  fin  is  larger  than 
the  second  dorsal  fin,  its  posterior  margin  is  only 
slightly  concave ;  the  pectorals  are  broadly  triangu- 
lar, their  anterior  margins  about  as  long  as  the 
distance  from  the  level  of  their  own  points  of 
origin  to  the  front  of  the  mouth. 

Color. — Gray  or  grayish  brown  above,  and  a 
paler  shade  of  the  same  below;  some  are  marked 
with  a  few  small  dark,  roundish  spots  along  the 
sides. 

Size. — This  shark  is  much  smaller  than  the 
hammerhead,  rarely  exceeding  5  feet  in  length;  it 
is  said  to  reach  6  feet. 

General  range. — Tropical-warm  temperate  At- 
lantic; from  southern  Brazd  to  North  Carolina, 
in  the  west,  and  as  a  stray  to  southern  New 
England  and  Massachusetts  Bay;  tropical  West 
Africa  in  the  east;  also  from  southern  California 


Figure  15. — Shovel  head  (Sphyrna  tiburo),  female,  about  14%  inches  long,  from  Rio  de  Janeiro.  A,  under  side  of  head; 
B,  first  to  seventh  upper  teeth  and  first  to  sixth  lower  teeth  counted  from  center  of  jaw,  about  3.6  times  natural  size. 
From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


FISHES    OF   THE   GULF   OF    MAINE 


45 


to  Ecuador  on  the  Pacific  Coast  of  America,  or 
represented  there  by  a  very  close  relative.9 

Occurrence  in  the  Gulf  oj  Maine. — Our  only 
reason  for  including  the  shovel-head  here  is  that 
a  stray  specimen  has  been  reported  from  Massa- 
chusetts Bay.10  It  has  also  been  taken  once  at 
Newport,  R.  I.,  and  a  commercial  shark  fishery 
that  was  carried  on  in  Nantucket  Sound  in  the 
summer  of  1918  is  said  to  have  yielded  six  of  them.11 

Common  hammerhead  Sphyrna  zygaena 
(Linnaeus)  1758 
Bigelow  and  Schroeder,  1948,  p.  436. 


'  On  this  point,  see  Bigelow  and  Schroeder,  Fishes  of  the  Western  North 
Atlantic,  Pt.  1,  1948,  p.  425,  footnote  20.  A  shark  has  also  been  reported  as 
tiburo  from  China  and  from  the  Philippines,  but  without  convincing  evidence 
as  to  its  identity. 

10  By  Oarman,  Mem.  Mus.  Comp.  Zool.,  vol.  36,  1913,  p.  161.  Apparently 
the  specimen  is  no  longer  in  existence. 

11  Personal  communication  by  R.  n.  Bodrnan,  who  operated  this  fishery. 


Description. — The  very  differently  shaped  head 
of  the  hammerhead,  the  shape  of  its  anal  fin  with 
much  more  deeply  concave  posterior  margin,  and 
the  fact  that  the  outermost  four  or  five  of  its  lower 
teeth  on  each  side  are  blade-like,  like  those  nearer 
the  center  of  its  mouth,  are  ready  field  marks  to 
separate  the  hammerhead  from  the  shovelhead 
(cf.  fig.  16  with  fig.  15).  The  anal  fin,  too,  is  only 
about  as  large  as  the  second  dorsal  in  the  hammer- 
head (considerably  larger  than  the  second  dorsal 
in  the  shovelhead).  Otherwise  the  positions  and 
shapes  of  the  fins  and  the  size  and  shape  of  the 
tail  are  much  alike  in  the  two  species. 

Color. — Leaden  or  brownish  gray  above,  shading 
along  the  sides  to  pure  or  grayish  white  below;  the 
tips  and  edges  of  the  dorsal  and  caudal  fins  are 
more  or  less  dusky;  and  the  tips  of  the  pectorals 
are  black  on  some  specimens. 


Figure  16. — Hammerhead  (Sphyrna  zygaena),  female,  about  27  inches  long,  from  Nahant,  Massachusetts.  A,  head 
from  below,  about  one-third  natural  size;  B,  second  upper  tooth;  C,  ninth  upper  tooth;  D,  third  lower  tooth; 
E,  ninth  lower  tooth;  about  4  times  natural  size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


46 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE    SERVICE 


Size. — It  appears  that  hammerheads  are  com- 
monly about  19  to  20  inches  long  when  they  are 
born;  seemingly,  they  mature  sexually  at  about  7 
to  8  feet;  they  are  often  taken  9  to  11  feet  long, 
and  occasionally  as  long  as  12  to  13  feet.12  Most  of 
those  that  visit  southern  New  England  are  less 
than  6  to  7  feet  long,  some  very  small  indeed.13 
In  1805,  however,  one  of  11  feet  was  netted  at 
Riverhead,  L.  I.  And  the  fact  that  it  contained 
parts  of  a  man  in  its  stomach  has  been  chiefly 
responsible  for  the  bad  reputation  of  this  species 
of  hammerhead. 

Two  other  large  sharks  closely  related  to  the 
common  hammerhead,  the  tropical  hammerhead 
(Sphyrna  lewini  Griffith,  1834) u  and  the  great 
hammerhead  (Sphyrna  mokarran  Ruppell,  1835)  16 
occur  along  the  South  Atlantic  coast  of  the  United 
States.  The  first  of  these,  in  particular,  might 
stray  as  far  as  Cape  Cod,  as  many  tropical  fishes 
do,  for  it  has  been  recorded  from  the  offing  of 
Cape  May,  New  Jersey.  They  resemble  the  com- 
mon hammerhead  closely  in  general  appearance, 
but  both  of  them  may  be  distinguished  from  the 
latter  by  the  fact  that  the  front  outline  of  their 
head  is  scalloped  in  the  midline,  not  evenly 
rounded  there  as  it  is  in  the  common  hammerhead. 
For  further  accounts  of  them,  see  Bigelow  and 
Schroeder.16 

Habits. — Since  hammerheads  are  an  accidental 
visitor  to  the  Gulf,  we  need  only  remark  that 
they  are  pelagic  in  habit,  often  swimming  with 
dorsal  and  caudal  fins  out  of  water,  and  are  to  be 
met  with  indifferently  out  at  sea  or  near  land. 
They  feed  chiefly  on  fish,  including  smaller 
sharks  (including  their  own  kind),  and  sting  rays, 

11  The  larger  hammerheads  that  are  sometimes  reported  probably  are  not 
this  species,  but  the  great  hammerhead  (Sphyrna  mokarran,  p.  46,  note  16). 

'»  Dozens  of  little  ones,  of  about  2H  feet,  have  been  seined  on  the  outer  shore 
of  Long  Island,  N.  Y.,  In  August. 

"  The  account  of  this  species,  In  Bigelow  and  Schroeder,  (Fishes  of  the 
Western  North  Atlantic,  Pt.  1,  1948,  p.  415)  was  as  diplana  Springer,  1941. 
But  Fraser-Brunner  (Rec.  Austral.  Mus.,  vol.  22,  No.  3,  1950,  pp.  213-214), 
has  shown  that  It  cannot  be  separated  from  the  Indo-Paclflc  S.  leuinl  of 
Griffith,  1834,  a  much  older  name. 

i*  Tortonese  has  recently  pointed  out  (Ann.  Mag.  Nat.  Hist.  Ser.  12,  vol.  3, 
No.  36,  1950,  p.  214)  that  the  name  tudes  Valenciennes  1822  that  has  been 
applied  commonly  to  the  great  hammerhead  of  the  Atlantic  actually  belongs 
to  a  different  species;  consequently  that  the  correct  name  of  the  great  hammer* 
head  Is  mokarran  RQppell,  1835,  It  being  Identical  with  that  Indo-Paclflc 
species. 

n  Fishes  Western  North  Atlantic,  Pt.  1, 1948,  pp.  415,  428. 


the  tail  spines  of  which  are  sometimes  found 
imbedded  in  their  jaws.  Like  tiger  sharks,  they 
make  themselves  a  pest  in  warmer  latitudes  where 
fisheries  for  sharks  are  carried  on,  by  devouring 
those  that  they  find  entangled  in  the  nets.  As 
many  as  30  to  37  embryos  have  been  found  in  a 
gravid  female,  and  the  embryos  do  not  develop  any 
placental  connection  with  the  mother,  so  far  as  is 
known. 

General  range. — Widespread  in  the  tropical  to 
warm  temperate  belts  of  the  Atlantic,  of  the 
Pacific,  and  probably  of  the  Indian  Ocean  as 
well;  north  commonly  to  southern  New  England, 
straying  to  Massachusetts  Bay  and  as  far  as 
Halifax,  Nova  Scotia.17 

Occurrence  in  the  Gulf  of  Maine. — Hammer- 
heads (often  in  small  schools)  wander  northward 
every  summer,  along  the  Atlantic  seaboard;  they 
are  often  to  be  seen  basking  at  the  surface  (some 
harpooned)  a  few  miles  out,  off  Marthas  Vine- 
yard and  Nantucket;  and  one  is  occasionally 
taken  in  one  or  another  of  the  fish  traps  near 
Woods  Hole.  But  the  longitude  of  Cape  Cod  so 
sharply  bounds  their  yearly  dispersal  that  the 
only  records  from  the  Gulf  of  Maine,  or  from 
Nova  Scotia  waters,  are  of  stray  specimens  from 
Chatham  and  Provincetown  on  the  outer  shores 
of  the  Cape;  of  one  about  27  inches  long  from 
Nahant,  in  the  inner  part  of  Massachusetts 
Bay ; I8  of  two  small  ones  recently  from  Casco 
Bay; 19  of  one  taken  many  years  ago,  off  Brier  I., 
on  the  Nova  Scotian  side  of  the  Bay  of  Fundy;20 
of  a  12-footer  harpooned  between  Georges  and 
Browns  Banks  in  August  1928  by  the  sword 
fishing  schooner  Doris  M.  Hawes;  of  a  small  one 
caught  in  Halifax  Harbor,  Nova  Scotia,  in 
September  1932;21  and  of  another  about  21  inches 
long  taken  in  a  trap  off  Sambro  Head,  near  Hali- 
fax, August  25,  1938.22 


"  For  further  details  of  distribution,  see  Bigelow  and  Schroeder,  Fishes  of 
the  Western  North  Atlantic,  Pt.  1,  1948,  p.  442. 

18  This  specimen,  obtained  many  years  ago  by  Louis  Agasslz,  Is  In  the 
Museum  of  Comparative  Zoology. 

'•  Seen  In  the  flsh  market  at  Portland,  Maine,  by  the  late  Walter  H.  Rich. 

»  McKenzle,  Proc.  Nova  Scotia  Inst.  Sci.,  vol.  20, 1939,  p.  13. 

"  Vladykov,  Proc.  Nova  Scotia  Inst.  Sci.,  vol.  19,  Pt.  1, 1935.  p.  8. 

"  McKenzle,  Prov.  Nova  Scotia  Inst.  Sci.,  vol.  20,  1939,  p.  13. 


FISHES   OF  THE  GULF  OF   MAINE 

THE  SPINY  DOGFISHES.     FAMILY  SQUALIDAE 


47 


This  group  is  characterized  by  the  lack  of  an 
anal  fin,  combined  with  the  presence  of  two  dorsal 
fins,  each  of  which  is  preceded  by  a  fixed  spine 
which  is  long  and  conspicuous  in  some,  but  so 
short  in  others  that  its  presence  can  be  detected 
only  by  touch.  The  teeth  are  alike  in  the  two 
jaws  in  some,  unlike  in  others. 

Spiny  dogfish  Squalus  acanthias  Linnaeus  1758 

Dogfish;  Piked  dogfish;  Grayfish 

Bigelow  and  Schroeder,  1948,  p.  455. 
Garman,  1913,  pi.  14,  figs.  1-4. 

Description. — Any  little  gray  or  brownish  shark, 
with  a  large  sharp  spine  lying  along  the  front 
margin  of  each  dorsal  fin,  caught  within  the  Gulf, 
or  on  the  shoaler  parts  of  the  offshore  fishing 
banks,  is  practically  sure  to  be  this  "dog,"  of 
which  there  are  thousands  in  the  Gulf  to  every 
one  shark  of  any  other  kind.  One  of  its  relatives, 
the  black  dogfish  (p.  51),  is  a  regular  inhabitant 
of  the  deeper  slopes  of  the  offshore  Banks  that 
front  the  Gulf,  where  we  also  trawled  more  than 
50  specimens  of  another  relative  Etmopterus 
princeps  Collett  1904  during  the  summer  of  1952. 
But  there  is  no  danger  of  confusing  the  common 
spiny-dog  with  either  of  these,  for  they  are  velvety 
black  in  color,  the  rear  margins  of  their  tail  fins 
are  indented  near  the  tip,  which  is  not  the  case  in 


the  spiny-dog,  and  each  of  their  teeth,  at  least  in 
the  upper  jaw  (lower  jaw  as  well  in  the  black  dog- 
fish) has  3  to  5  sharp  points,  but  only  one  point 
in  the  spiny  dog. 

This  is  a  slender  little  shark,  with  flattened 
head  and  snout  tapering  to  a  blunt  tip.  Its  first 
dorsal  fin  stands  between  pectorals  and  pelvics; 
its  second  dorsal  fin  is  about  two-thirds  as  large 
as  the  first;  its  pectorals  form  nearly  an  equilateral 
triangle;  and  its  pelvics  are  well  forward  of  its 
second  dorsal  fin.  The  dorsal  fin  spines  he  close 
along  the  front  margins  of  the  two  dorsals,  the 
first  not  more  than  one-half  as  long,  and  the  second 
nearly  as  long  as  the  front  margin  of  their  respec- 
tive fin,  and  they  are  very  sharp.  The  spiny-dog 
has  no  anal  fin,  a  lack  separating  it  from  all 
smooth-finned  sharks  known  from  the  Gulf  of 
Maine,  except  for  the  Greenland  shark  (p.  53), 
Dalatias  (p.  55),  and  the  bramble  shark  (p.  56). 
There  is  a  low  fold  of  skin  on  either  side  of  the 
root  of  the  tail  back  of  the  second  dorsal  fin,  so 
small,  however,  that  there  is  no  danger  of  confusing 
it  with  the  caudal  keels  of  the  mackerel-shark 
tribe.  The  teeth  are  small,  their  sharp  points 
bent  toward  the  outer  corners  of  the  mouth  so 
that  they  form  a  nearly  continuous  cutting  edge 
along  each  jaw. 

Color. — The  upper  surface  is  slate  colored  usu- 
ally, sometimes  tinged  brown,  with  a  row  of  small 


Fiqube  17. — Spiny  dogfish  (Squalus  acanthias),  female,  27  inches  long;  after  Garman.  A,  upper  and  lower  teeth,  mid- 
point of  mouth  marked  by  the  dotted  line,  about  3  times  natural  size.  From  Bigelow  and  Schroeder.  Drawing 
by  E.  N.  Fischer. 


48 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


white  spots  on  each  side  from  the  pectoral  fin  to 
abreast  of  the  anal  fin,  and  with  a  few  other 
white  spots  in  front  of  the  first  dorsal  and  behind 
it,  also  in  front  of  the  second  dorsal  fin.  These 
spots  are  most  conspicuous  on  small  fish  up  to  12 
or  14  inches  long  and  they  fade  with  growth  until 
they  disappear  altogether  in  some  specimens. 
The  margins  of  the  first  and  second  dorsals,  and  of 
the  caudal  are  more  or  less  dusky  at  birth,  but 
soon  fade.  The  lower  surface  ranges  from  pale 
gray  to  pure  white. 

Size. — The  majority  are  between  8%  and  13 
inches  long  when  born.  Most  of  the  adult  males 
are  from  about  2  feet  to  a  little  less  than  3  feet 
long;  adult  females  are  from  a  little  less  than  2% 
feet  to  almost  3%  feet;  maximum  length  about 
four  feet.  Mature  females  average  7  to  10  pounds, 
a  few  reach  15  pounds  if  very  fat,  and  20  pounds 
has  been  reported. 

Habits. — Much  has  been  written  of  the  habits 
of  the  spiny  dogfish,  but  nothing  to  recommend 
it  from  the  standpoint  either  of  the  fishermen  or 
of  its  fellow  creatures  in  the  sea.  It  is  one  of  the 
more  gregarious  of  our  fishes,  swimming  in  schools 
or  packs.  Swedish  fishermen  assert  that  young 
dogs  school  separately  from  their  parents,  and  it 
is  certain  that  fish  of  a  size  continue  to  associate 
together  as  they  grow,  the  result  being  that  any 
given  school  runs  very  even,  consisting  as  a  rule 
either  of  the  very  large  mature  females,  or  of 
medium-sized  fish  (either  mature  males  or  im- 
mature females) ,  or  of  small  immature  fish  of  both 
sexes  in  about  equal  numbers. 

Apart  from  their  general  seasonal  migratory 
movements,  dogfish  are  governed  by  the  move- 
ments of  the  fishes  on  which  they  prey.  And  re- 
cent marking  experiments  have  shown  that  some 
of  them  cover  long  distances  in  their  wanderings, 
for  two  tagged  near  St.  Johns,  Newfoundland,  in 
mid-July  1942  were  recaught  off  Cape  Ann,23 
one  on  November  23,  1943,  the  other  on  Decem- 
ber 4  of  that  year,24  while  others  from  the  same 
tagging  experiment  were  caught  within  the  Gulf 
of  St.  Lawrence.25  Fortunately  they  seldom  stay 
long  in  one  place,  but  there  is  seldom,  if  ever,  a 
time  during  the  summer  when  they  are  not  com- 
mon on  some  part  of  the  Gulf  of  Maine  coast. 
So  erratic  are  their  appearances  and  disappearances 

23  About  14  miles  offshore. 

24  On  Middle  Ground  about  25  miles  off  Cape  Ann. 

"  Templeman,  Fish.  Res.  Bull.,  Newfoundland  Dept.  Nat.  Res.,  No.  15, 
1944,  pp.  67-69. 


that  where  one  has  good  fishing  today  he  may 
catch  only  dogfish  tomorrow  and  nothing  at  all 
the  day  after,  the  better  fish  having  fled  these  sea 
wolves  and  the  latter  departing  in  pursuit. 

The  dogfish  use  their  back  spines  for  defense, 
curling  around  in  a  bow  and  striking,  which  makes 
them  hard  to  handle  on  the  hook.  It  is  probable, 
too,  that  the  spines  are  slightly  poisonous,  general 
report  to  this  effect  being  corroborated  by  the 
fact  that  the  concave  surfaces  are  lined  with  a 
glandular  tissue  resembling  the  poison  glands  of 
the  venomous  "weever"  (Trachinus  draco)  26  of 
Europe. 

Voracious  almost  beyond  belief,  the  dogfish 
entirely  deserves  its  bad  reputation.  Not  only 
does  it  harry  and  drive  off  mackerel,  herring,  and 
even  fish  as  large  as  cod  and  haddock,  but  it 
destroys  vast  numbers  of  them.  Again  and  again 
fishermen  have  described  packs  of  dogs  dashing 
among  schools  of  mackerel,  and  even  attacking 
them  within  the  seines,  biting  through  the  net, 
and  releasing  such  of  the  catch  as  escapes  them. 
At  one  time  or  another  they  prey  on  practically  all 
species  of  Gulf  of  Maine  fish  smaller  than  them- 
selves, and  squid  are  also  a  regular  article  of  diet 
whenever  they  are  found.  Dogfish  are  also  known 
to  take  worms,  shrimps,  and  crabs.  And  when 
they  first  arrive  at  Woods  Hole  in  May  they 
are  often  found  full  of  Ctenophores,  being  one 
of  the  few  fish  that  eat  these  watery  organisms. 
Often,  too,  they  bite  groundfish  from  the  hooks 
of  long  lines,  or  take  the  baits  and  make  it  futile 
to  fish  with  hook  and  line  where  they  abound. 

Fishermen  are  familiar  with  the  fact  that  the 
female  spiny  dog  bears  "living"  young  (this  has 
been  known  since  the  days  of  Aristotle).  The 
eggs  are  large,  well  stored  with  yolk,  and  during 
early  stages  those  in  each  oviduct  (so-called 
"uterus")  are  contained  in  a  horny  capsule  that 
breaks  down  later,  leaving  the  embryos  free  in  the 
"uterus,"  to  which  they  have  no  placental  attach- 
ment. The  number  in  a  litter  is  commonly  4  to  6; 
sometimes  as  many  as  8  to  11,  or  as  few  as  2. 

According  to  recent  studies,  the  females  carry 
their  young  for  18  to  22  months.  Accordingly,  the 
adult  females  caught  in  our  Gulf  contain  either 
very  early  embryos,  averaging  only  about  three- 
fourths  of  an  inch  in  length  by  September,  or 


*  Evans  (Pbilos.  Trans.  Royal  Soc.,  London,  Ser.  B,  vol.  212,  1923,  pp.  8, 
27)  describes  the  spines  and  gives  clinical  records  of  the  effects  of  wounds 
inflicted  by  them. 


FISHES    OF   THE   GULF   OF    MAINE 


49 


much  larger  ones,  7  to  11  inches  long  by  that 
month;  i.  e.,  nearly  ready  for  birth.  Similarly, 
we  have  taken  females  with  embryos  9  to  10% 
inches  long  in  November,  on  the  Cholera  Bank 
near  New  York  Harbor.  And  it  now  seems 
established  that  most  of  the  young  are  born  on  the 
offshore  wintering  grounds.27  But  dogfish  so 
small  as  evidently  to  have  been  newborn  are  oc- 
casionally taken  along  southern  New  England 
and  in  the  Gulf  in  early  summer;  also  on  Nantucket 
Shoals  where  the  Albatross  II  trawled  some  of 
10}£  to  13  inches  in  August,  showing  that  the 
season  of  production  extends  through  the  spring,  or 
even  into  the  summer  as  in  1905  when  females 
taken  off  Gloucester  in  July  gave  birth  to  young 
on  capture.28 

General  range. — Both  sides  of  the  North  Atlantic, 
chiefly  in  the  temperate  and  subarctic  belt;  also 
both  sides  of  the  northern  Pacific; 29  and  repre- 
sented in  the  corresponding  thermal  belt  of  the 
southern  hemisphere  by  a  relative  (or  relatives) 
so  close  that  it  is  doubtful  whether  they  differ  in 
any  recognizable  way  from  the  spiny -dog  of  the 
north. 

Occurrence  in  the  Gulf  of  Maine.  The  spiny  dog- 
fish ("dogfish"  or  "dog"  in  common  parlance)  makes 
up  for  the  comparative  rarity  of  other  sharks  in 
the  Gulf  of  Maine  by  its  obnoxious  abundance. 
To  mention  all  the  localities  from  which  it  has  been 
reported  there  would  be  simply  to  list  everv  seaside 
village  and  fishing  ground  from  Cape  Cod  to  Cape 
Sable.  It  is  as  familiar,  too,  on  the  offshore  banks 
as  it  is  along  the  coast;  also  along  outer  Nova 
Scotia,  in  the  Gulf  of  St.  Lawrence,  on  the  Grand 
Banks,  and  along  the  east  coast  of  Newfoundland 
to  southeastern  Labrador.  There  is  no  record  of 
it  from  the  North  American  coast  north  of  Hamil- 
ton Inlet,  but  stray  specimens  have  been  taken 
along  the  southwest  coast  of  Greenland.30  To  the 
southward,  fishermen  are  familiar  with  it  in  season 

"  Females  that  we  saw  trawled  oft  Block  Island  in  (50-65  fathoms  in  late 
January  1950,  gave  birth  to  young  on  the  deck  of  the  vessel. 

»  Mclntire,  Rept.  Comm.  Fish.  Game  Massachusetts,  (1905)  1906,  p.  108. 

»  We  have  found  no  consistent  differences  between  North  Atlantic  and 
North  Pacific  specimens.  For  further  discussion  of  this  point,  and  further 
details  as  to  the  occurrence  of  the  spiny-dog  in  the  two  sides  of  the  North  At- 
lantic, see  Bigelow  and  Schroeder  (Fishes  of  the  Western  North  Atlantic, 
Pt.  1,  1948,  pp.  453,  463). 

"Jensen  (Selachians  of  Greenland,  Mindeskr.  Japetus  Stcenstrup,  Pt.  2, 
No.  30, 1914,  p.  7)  lists  several  definite  records  of  this  species  at  Sukkertoppen 
and  near  nolsteinborg.  West  Greenland. 


as  far  as  Cape  Lookout,  N.  C,  and  a  few  stray 
even  to  southern  Florida  and  to  Cuba.31 

Dogfish  are  seasonal  visitors  on  the  coast,  strik- 
ing in  about  as  early  along  New  Jersey  (March), 
and  even  on  Georges  Bank  (March-April),  as 
along  North  Carolina.  In  the  inner  parts  of  the 
Gulf  of  Maine  the  date  of  the  first  heavy  run  of 
dogfish  varies  widely  from  year  to  year  and  from 
place  to  place.  We  have  not  heard  of  them  there 
before  May.  But  the  period  of  freedom  may  close 
as  early  as  the  last  half  of  the  month,  in  some  years. 

In  1903,  for  example,  they  had  appeared  as  far 
north  as  Penobscot  Bay  by  the  middle  of  May. 
And  while  it  is  not  until  June  that  they  usually 
arrive  in  numbers  in  the  Massachusetts  Bay  re- 
gion, it  is  sometimes  impossible  to  set  gill  or  drift 
nets  anywhere  between  Cape  Cod  and  Cape 
Elizabeth  after  the  first  days  of  that  month,  so 
numerous  are  they.  In  1913  the  first  heavy  run 
of  dogfish  struck  Ipswich  Bay  on  June  14,  and  they 
appeared  there  at  about  the  same  date  in  1905, 
but  there  is  much  local  variation  in  this  respect. 
In  1903,  for  example,  they  did  not  appear  until 
early  July  at  Province  town,  though  swarming  a 
month  earlier  in  Massachusetts  Bay,  in  Ipswich 
Bay,  and  off  Penobscot  Bay.  But  in  1920  they 
appeared  at  Provincetown  by  May  25  to  26  when 
one  set  of  mackerel  traps  caught  23  barrels  of  them, 
and  another  21  barrels.  They  usually  strike  in 
all  along  the  northern  Maine  and  west  Nova 
Scotia  coasts  by  the  end  of  June;  but  few  are  seen 
until  late  in  July  in  Passamoquoddy  Bay.  They 
have  been  recorded  as  early  as  July  1  near  Raleigh, 
on  the  Newfoundland  side  of  the  Strait  of  Belle 
Isle,  but  they  are  not  caught  in  any  numbers  in 
the  inner  parts  of  the  Gulf  of  St.  Lawrence  until 
well  into  July,  and  they  have  not  been  reported 
from  southeastern  Labrador  until  early  in  Sep- 
tember.32 

In  the  southern  part  of  its  range,  from  North 
Carolina  to  New  York,  the  spiny  dogfish  is  a  spring 
and  autumn  transient  only.  West  of  Cape  Cod 
(at  Woods  Hole,  that  is,  and  along  Long  Island) 

"  Repeated  reports  of  it  as  plentiful  along  eastern  Florida  seem  to  have 
referred  to  some  other  shark;  the  basis  for  similar  reports  from  Cuba  and 
Trinidad  doubtless  was  the  Cuban  dogfish,  Squalm  cubensis  Rivero. 

"  See  Templeman  (Res.  Bull.  15,  Newfoundland  Dept.  Nat.  Res.,  1941, 
pp.  56,  64)  for  dates  of  arrival  around  the  coast  of  Newfoundland  in  different 
years. 


50 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


they  are  transients  mostly,  passing  north  in  spring 
and  south  in  autumn,  though  some  summer  there; 
even  considerable  numbers  in  some  years.33  And 
it  seems  that  most  of  them  withdraw  from  Mas- 
sachusetts Bay  also  during  the  warmest  period, 
for  few  are  taken  there  between  June  and  Septem- 
ber. But  they  continue  present  all  summer  along 
outer  Cape  Cod,  and  here  and  there  throughout 
the  northern  and  eastern  parts  of  the  Gulf,  in 
varying  abundance. 

Most  of  the  dogfish  take  their  autumnal  depar- 
ture from  the  inner  parts  of  the  Gulf  during 
October,  few  being  caught  on  the  coast  north  of 
Massachusetts  Bay  after  November  1.  But  they 
sometimes  stay  later,  as  in  1903  (a  big  dogfish 
year),  and  again  in  1942,  when  they  were  abundant 
along  the  outer  shore  of  Cape  Cod  as  late  as  the 
first  week  of  November.  Ordinarily  none  are 
caught  within  the  Gulf  of  Maine  north  of  Georges 
Bank  in  winter,  but  this  has  its  exceptions.  In 
1913,  for  example,  a  few  were  caught  20  miles  off 
Cape  Ann  on  November  19  to  24,  many  near 
Boon  Island  from  December  5  to  13,  and  on 
Jeffreys  Ledge  on  December  11  and  12. 

In  1882,  schools  were  reported  off  Portsmouth, 
N.  H.,  even  as  late  as  February,  an  exceptional 
event. 

Dogfish  appear  earlier  in  spring  and  linger 
later  into  the  winter  on  Georges  Bank  (fig.  18) 
than  in  the  inner  parts  of  the  Gulf.  It  is  safe  to 
say  that  there  are  few  there  in  March,  the  earliest 
definite  record  (obtained  during  the  investigations 
of  1913,  only  year  of  record,  being  of  25  fish  caught 
on  the  "winter  cod  ground"  east  of  the  shoals 
(long,  about  67°,  lat.  about  41°40')  between  the 
20th  and  the  22nd,  and  of  46  from  the  same  gen- 
eral region  from  the  27th  to  the  30th,  while  some 
are  trawled  there  all  summer.  In  1913,  a  few 
were  taken  in  November  and  in  December;  a 
few  also  on  the  southern  part  of  the  Bank  (lat. 
about  41°,  long,  about  67°30')  on  January  20  to 
22  in  1914. 

Apparently  dogfish  reach  Browns  Bank  later 
than  they  do  Georges,  for  none  was  taken  there 
on  April  14  in  1913,  though  they  are  only  too 
plentiful  there  in  summer.  It  is  also  likely  that 
they  depart  earlier,  although  a  few  lingered  as  late 
as  December  3  to  12  on  Western  Bank  off  Halifax 
in  that  year. 

»  For  details,  see  Bigelow  and  Schroeder,  Fishes  of  the  Western  North 
Atlantic.  Ft.  1,  1948,  p.  464. 


ouuu 
7500 

7000 

6500 

eooo 

|    5500 
2    5000 

g    4500 

5    4000 

I    3500 

£    3000 

Si 

f    2500 

K 

2000 

• 

1500 

■ 

1000 

■ 

500 

..     .     I 

I 

■    I. 

ll 

1 

MAR.  'Will  '  MAY 

JUNE' JULY 

' 

AUG. 

SEPT.  '    OCT. '    NOV. ' 

DEC. 

Figure  18. — Numbers  of  spiny  dogfish  caught  on  certain 
otter  trawling  trips  to  Georges  Bank,  during  the  dif- 
ferent months  of  1913. 

It  now  seems  certain  that  the  spiny  dogfish 
winter  chiefly  in  deeper  water  offshore,  for  con- 
siderable numbers  have  been  trawled  at  that 
season  on  the  outer  part  of  the  continental  shelf 
off  Block  Island,  in  50  to  65  fathoms,  where  we 
saw  several  hundred  (200  in  one  haul)  trawled 
during  the  last  week  of  January  1950;  off  New 
York  in  November  and  January; 34  also  in  Febru- 
ary off  the  Middle  Atlantic  coast  in  16  to  70 
fathoms,  south  as  far  as  the  offing  of  Cape  Hat- 
teras.  On  the  other  hand,  the  fact  that  numbers 
of  them  have  been  found  washed  on  shore  in 
January  on  the  southwest  coast  of  Newfoundland 
suggests  that  some  of  those  that  summer  in  that 
general  region  may  survive  the  winter  in  the  deep 
trough  of  the  Gulf  of  St.  Lawrence.  They  are 
usually  so  thin  when  they  reappear  on  the  coast 
in  spring  as  to  suggest  that  they  feed  but  little 
during  the  winter. 

This  is  the  only  Gulf  of  Maine  shark  that  even 
remotely  rivals  the  important  food  fishes  in  num- 
bers. Unfortunately,  the  statistics  of  the  com- 
mercial landings  for  American  waters  do  not 
afford  any  information  in  this  regard.  But  spiny 
dogs  must  be  plentiful  indeed  in  our  waters  when 
they  can  sometimes  be  caught  as  fast  as  they  can 

u  Mr.  Thomas  Quast  informs  us  that  many  were  taken  from  the  schooner 
Victor,  long-lining  for  tile  fish,  on  the  outer  edge  of  the  continental  shelf,  off 
New  York,  during  the  second  week  of  January  1928. 


FISHES    OF   THE   GULF   OF    MAINE 


51 


be  hauled  in;  when  a  long  line,  with  1,500  hooks, 
has  been  known  to  bring  in  a  dogfish  on  nearly 
every  hook ;  and  when  an  average  trawl  catch  of 
6,000  to  8,000  per  trip  was  made  on  Georges  Bank 
in  1913  during  their  season  of  abundance.  At 
the  time  of  the  1904  to  1905  peak  it  was  estimated 
from  recorded  catches  that  at  least  27,000,000 
were  being  taken  yearly  off  the  coast  of  Massachu- 
setts.35 

More  precise  information  from  waters  farther 
north  is  that  10,391,000  pounds,  or  2  to  3  million 
individual  dogfish,  were  caught  in  1938,  in  Pla- 
centia  Bay,  Newfoundland,  with  no  apparent 
effect  on  their  numbers.36  In  short,  they  may  be 
as  plentiful  in  our  Gulf  as  they  are  on  the  Cornish 
coast,  where  the  record  catch  of  20,000  in  a  single 
haul  was  made  many  years  ago. 

Spiny  dogfish  appear  to  have  been  more  numer- 
ous in  the  Massachusetts  Bay  region  during  the 
last  quarter  of  the  past  century  and  during  the 
early  nineteen  hundreds  than  they  had  been  pre- 
viously. At  Woods  Hole,  on  the  contrary,  they 
are  said  to  have  been  much  more  plentiful  before 
1887  than  they  have  been  at  any  time  since.  To 
a  certain  extent,  of  course,  reports  of  fluctuations 
in  abundance  from  year  to  year  must  be  discounted 
as  reflecting  the  movements  of  the  great  schools 
that  may  visit  one  part  of  the  coast  one  summer 
and  another  part  the  next,  not  a  general  altera- 
tion of  the  stock.  But  the  many  fishermen  who 
reported  to  the  Massachusetts  Commissioners  in 
1905  were  unanimously  of  the  opinion  that  dogfish 
had  multiplied  steadily  for  20  to  30  years  past, 
and  reports  from  British  coasts  were  to  the  same 
effect.  Perhaps  the  years  1904-1905  marked  the 
apex  of  this  wave  of  multiplication;  at  any  rate 
dogfish  were  reported  as  distinctly  less  troublesome 
to  the  mackerel  netters  in  1913  than  they  had 
been  previously.  And  little  complaint  has  been 
made  of  them  in  late  years. 

But  it  is  not  safe  to  conclude  from  this  that  the 
stock  is  at  a  low  ebb  at  present,  for  it  was  the  hand- 
and  long-line  fishermen  that  suffered  most  from 
them;  and  it  is  only  as  they  increase  the  amounts 
of  trash  fish  dumped  overboard  that  the  dogfish 
bother  the  otter-trawlers. 

Importance. — During  the  years  when  the  ground 
fishery  was  chiefly  by  hook  and  line,  fishing  often 
was  actually  prevented  by  dogfish  in  Massachu- 

»  Report,  Comm.  Fish  and  Game,  Mass.,  (1906),  1907,  p.  20. 
»  Templeman,  Newfoundland  Fish.    Res.  Bull.,  15,  1944,  p.  72. 


setts  and  Ipswich  Bays,  unless  cockles  (Polynices) 
were  used  for  bait,  for  dogfish  do  not  take  these. 
The  general  replacement  of  hook  and  line  fishing 
by  the  otter  trawl  has  put  an  end  to  widespread 
complaints  on  this  score.  But  when  schools  of 
dogfish  get  into  a  net  or  seine,  they  so  snarl  the 
twine  that  disentanglement  and  repair  may  be  the 
work  of  days.  And  it  has  been  estimated  that 
they  may  do  some  $400,000  worth  of  damage 
annually  to  fishing  gear,  and  to  fish  caught  by 
such  gear,  off  the  coast  of  Massachusetts  alone, 
during  their  peaks  of  abundance  there. 

With  the  dogfish  so  plentiful  and  destructive, 
it  is  no  wonder  that  serious  efforts  have  been  made 
to  make  them  a  source  of  revenue  instead  of  a 
dead  loss.  And  the  dog  is  a  far  better  food  fish 
when  fresh  than  is  generally  appreciated,  as  is 
evident  by  the  large  amounts  landed  in  the  fishing 
ports  of  northwestern  Europe.  But  it  has  never 
been  in  any  demand  for  the  table,  on  our  coasts, 
though  it  would  offer  a  large  supply  of  cheap  food 
were  a  satisfactory  method  found  for  canning  it. 
During  their  more  recent  periods  of  plenty  various 
efforts  have  been  made  to  utilize  them  on  a  large 
scale  for  fertilizer,  and  for  liver  oil  (it  compares 
favorably  with  cod  for  vitamin  A,  though  it  is 
much  poorer  in  vitamin  D) ,  on  the  Atlantic  coasts 
of  the  United  States  and  Canada;  however  such 
developments  have  been  short-lived.  And  dogfish 
have  not  been  of  sufficient  value  up  to  the  present 
to  compensate  for  a  hundredth  part  of  the  damage 
they  do.37 

Black  dogfish  Centroscyllium  fabricii 
(Reinhardt)  1825 

Bigelow  and  Schroeder,  1948,  p.  482. 
Garman,  1913,  pi.  10,  figs.  5-8. 

Description. — The  notched  margin  of  the  upper 
tail  lobe  distinguishes  this  shark  at  a  glance  from 
the  spiny  dogfish,  with  which  it  agrees  in  having  a 
long  pointed  spine  at  the  front  edge  of  each  dorsal 
fin.  It  differs  further  from  the  common  dogfish 
in  that  its  dorsal  spines  are  deeply  grooved  along 
each  side,  whereas  in  the  "dog"  they  are  rounded; 
in  the  location  of  the  pelvic  fins,  the  rear  axils  of 


87  For  further  discussion  of  the  damage  done  by  dogfish  and  of  their  com- 
mercial possibilities,  see  Ann.  Rept.,  Comm.  Fish.  Oame  Mass.  (1905),  1908, 
pp.  97-169;  Rept.  U.  S.  Comm.  Fish.  (1902)  1904,  pp.  228-229;  Field,  Doc. 
622,  Rept.  U.  S.  Comm.  Fish.  (1906)  1907,  pp.  21-23;  Field,  Bull.  U.  S.  Bur. 
Fish.,  vol.  28,  1910,  pp.  243-257;  Mayor,  Contr.  Canad.  Biol.  (1918-1920) 
1921,  pp.  125-135;  and  Templeman,  Newfoundland  Fish  Res.  Bull.  15,  1944 


52 


FISHERY    BULLETIN    OF    THE    FISH    AND   WILDLIFE    SERVICE 


Figure  19. — Black  dogfish  (Cenlroscyllium  fabricii) ,  female,  about  25  inches  long,  from  the  southeast  slope  of  Georges 
Bank.  A,  first  three  upper  teeth  counted  from  center  of  jaw;  B,  twentieth  upper  tooth;  C,  first  three  lower  teeth; 
D,  lower  sixteenth  tooth;  about  5  times  natural  size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


which  stand  almost  directly  under  the  front  origin 
of  the  second  dorsal  fin  instead  of  some  distance 
in  front  of  the  latter;  in  its  small  pectorals  of 
rounded  outline;  in  the  shapes  of  its  teeth,  each  of 
which  has  3  or  5  sharp  points;  in  its  broad  rounded 
snout;  and  in  its  very  dark  color.  Like  the  spiny 
dogfish,  it  lacks  an  anal  fin. 

Size. — Adult  specimens  range  from  2  to  3%  feet 
in  length,  that  is,  about  the  same  size  as  the  spiny 
dogfish. 

Color. — Uniform  dark  brown  to  black,  below  as 
well  as  above. 

Habits. — In  West  Greenland  waters  cephalopods, 
pelagic  crustaceans,  and  medusae  have  been  found 
in  their  stomachs,  and  females  have  been  taken 
with  embryos  in  February.  Perhaps  they  are 
luminescent,  for  their  skins  bear  minute  deeply 
pigmented  dots,  suggesting  the  light  organs  of 
the  brilliantly  luminescent  shark  Isistius  brasili- 
ensis. 

General  range. — Northern  North  Atlantic;  Faroe 
Bank,  Faroe-Shetland  Channel  and  Iceland  in  the 
east;  West  Greenland;  Davis  Strait;  and  outer 
slopes  of  the  fishing  banks  in  the  west,  southward 
to  Georges  Bank;  chiefly  deeper  than  150  fathoms. 

Occurrence  in  the  Gulf  of  Maine.  In  the  years 
when  a  long  line  fishery  for  halibut  was  carried  on 
regularly,  black  dogfish  were  often  caught  along 
the  slopes  of  the  offshore  Banks,  from  Grand  to 
Browns  and  to  the  eastern  part  of  Georges,  if 
sets  weremadedown  to  200  fathoms  or  deeper.  And 
while  they  dropped  out  of  sight  with  the  general 
abandonment  of  that  fishery,  no  doubt  they  are  as 
plentiful  now  as  formerly,  for  we  trawled  about 
100  of  them,  6  to  24%  inches  long,  off  southwestern 
Nova  Scotia,  at  290  to  580  fathoms,  on  the  Caryn 


of  the  Woods  Hole  Oceanographic  Institution,  in 
June  1949.  How  far  they  range  to  the  west  and 
south,  at  the  appropriate  depths,  is  not  known.38 

Portuguese  shark  Centroscymnus  coelolepis 
Bocage  and  Brito  Capello,  1864 

Bigelow  and  Schroeder,  1948,  p.  494. 
Garman,  1913,  pi.  14,  figs.  5-8. 

Description. — This  shark  can  be  identified  easily 
by  the  fact  that  while  its  general  appearance 
(especially  the  absence  of  anal  fin,  the  situation  of 
its  pelvics  far  back  under  the  second  dorsal,  and 
its  rather  stout  form  and  blunt  snout)  might  lead 
a  hasty  observer  to  think  he  had  caught  a  small 
Greenland  shark;  more  careful  examination,  by 
touch  if  not  by  eye,  would  reveal  a  short  spine 
close  in  front  of  each  dorsal  fin.  The  first  dorsal 
fin  is  smaller  than  in  any  of  our  sharks  except  in 
the  "Greenland,"  (p.  53),  and  in  Dalatias  licha 
(p.  55),  the  second  dorsal  is  a  little  larger  than  the 
first,  and  the  pelvics  are  larger  than  either  of  the 
dorsals.  The  tail  is  noticeably  short  and  broad 
and  the  rear  edge  of  its  upper  lobe  is  notched.  The 
teeth  are  different  in  the  two  jaws;  narrow, pointed, 
and  of  the  seizing  type  in  the  upper;  broader,  ob- 
long, with  a  notch  on  the  outer  side  near  the  tip, 
and  forming  a  continuous  cutting  edge  in  the  lower. 
The  dermal  denticles  are  flat,  scale-like,  closely 
overlapping,  and  clothe  the  entire  trunk. 

Color. — Dark  chocolate  brown,  belly  as  well  as 
back  and  fins. 


38  Its  range  has  been  said  to  extend  to  New  York,  but  without  supporting 
evidence;  and  report  of  a  young  one  from  the  Gulf  of  Mexico  (Goode  and 
Bean,  Smithsonian  Contrib.  Knowledge,  vol.  30,  1895,  p.  11),  probably  was 
based  on  some  other  shark. 


FISHES    OF  THE   GULF   OF    MAINE 


53 


Figure  20. — Portuguese  shark  (Cenlroscymnus  coelolepis),  female  about  42%  inches  long,  off  Banquereau  Bank.  A, 
upper  teeth,  and  B,  lower  teeth  from  center  of  mouth,  about  3.4  times  natural  size.  From  Bigelow  and  Schroeder. 
Drawings  by  E.  N.  Fischer. 


Size. — Adults  measure  from  3  to  3%  feet  long, 
as  they  are  caught.  Garman  records  one  44 
inches  long  taken  off  the  coast  of  New  England. 
About  9  inches  is  the  smallest  recorded.39 

Habits. — Little  is  known  of  its  habits  beyond 
the  fact  that  it  is  a  deep-water  species,  and  that  it 
was  caught  regularly  by  Portuguese  fishermen 
with  hand  lines,  a  fishery  that  Wright 40  described 
as  follows: 

Some  600  fathoms  of  rope  were  let  out,  the  first  30  or 
40  fathoms  of  which  had  fastened  to  it  at  intervals  of  a 
fathom  a  series  of  small  ropes,  on  each  of  which  was  a  large 
hook  baited  with  a  codling.  This  fishing  tackle  remained 
below  for  about  two  hours,  when  they  commenced  to  haul 
it  in.  When  it  arrived  at  the  last  few  fathoms,  they  pulled 
in,  one  after  another,  five  or  six  specimens  from  3  to  4  feet 
long.     The    species    was    the     Cenlroscymnus    coelolepis 


Bocage  and  Capello.     These  sharks,  as  they  were  hauled 
into  the  boat,  fell  down  into  it  like  so  many  dead  pigs. 

Thirteen  to  16  young  have  been  found  in  fe- 
males caught  off  Portugal. 

General  range. — This  deep-water  shark,  origi- 
nally discovered  off  Portugal,  has  since  been  taken 
at  various  other  eastern  Atlantic  localities.41  Defi- 
nite records  of  it  for  the  western  Atlantic  are  from 
the  slopes  of  the  Nova  Scotian  Banks  and  of 
Georges,  at  depths  of  180  to  250  fathoms,  perhaps 
15  to  20  specimens  in  all.  But  Goode  and  Bean's  " 
old  characterization  of  them  as  abundant  on  the 
Banks  at  200  fathoms  and  deeper  presents  its  local 
status  more  correctly,  for  fishermen  long  lining 
for  halibut  often  caught  one  or  two  a  trip  in  the 
deeper  gullies  between  the  offshore  Banks. 


THE  GURRY  SHARKS.     FAMILY  DALATIIDAE 


The  gurry  sharks,  like  the  spiny  dogfishes,  lack 
anal  fins,  but  they  have  no  spines  in  their  dorsal 
fins.  The  teeth  in  the  upper  jaw  are  noticeably 
unlike  those  in  the  lower. 

Greenland  shark  Somniosus  microcephalus  (Bloch 
and  Schneider)  1801 

Sleeper  shark;  Gurry  shark;  Ground  shark 

Bigelow  and  Schroeder,  1948,  p.  516. 
Garman,  1913,  pi.  15,  figs.  4-6. 

Description. — The  Greenland  shark  is  notable 


»•  A  male  228  mm.  long,  examined  by  us,  in  the  U.  S.  National  Museum, 
from  the  continental  edge  south  of  Nantucket. 
••  Ann.  Mag.  Nat.  Hist.,  Ser.  4,  vol.  2, 1868,  p.  426. 


for  its  small  dorsal  fins,  without  spines,  the  second 
dorsal  being  of  about  the  same  size  as  the  first, 
and  for  small  pectorals  hardly  larger  than  the 
pelvics,  coupled  with  the  absence  of  an  anal  fin 
and  with  a  tail  of  more  fish-like  form  than  that  of 
most  other  sharks  except  for  the  mackerel-shark 
tribe.  Bearing  these  points  in  mind,  particularly 
the  absence  of  an  anal  fin  and  of  dorsal  spines,  it 
cannot  be  confused  with  any  shark  common  in 
our  Gulf.  And  while  it  resembles  the  rare 
Portuguese  shark  in  the  sizes  and  relative  situa- 


»  Iceland;  Faroe  Bank;  Madeira;  Azores;  Morocco;  Cape  Verde  I.:  For  key 
to  other  species  of  the  genus,  see  Bigelow  and  Schroeder,  Fishes  Western 
North  Atlantic,  P.  1,  1948,  p.  494. 

•>  Smithsonian  Contrib.  Knowledge,  vol.  30,  1895,  p.  14. 


54 


FISHERY   BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


Figure  21. — Greenland  shark  (Somniosus  microcephalus) , 
female,  about  5  feet  9  inches  long.  Teeth  at  center  of 
mouth ;  lower  teeth  from  midway  along  the  jaw  of  a  speci- 
men about  11  feet  long  from  the  Gulf  of  Maine,  about 
1.8  times  natural  size.  From  Bigelow  and  Schroeder. 
Drawings  by  E.  N.  Fischer. 


tions  of  its  dorsal  and  anal  fins,  in  its  general  form, 
and  in  its  teeth,  it  is  easily  separable  from  the 
"Portuguese,"  both  by  lacking  any  trace  of  spines 
in  its  dorsal  fins,  by  its  thorn-like  and  loosely 
spaced  dermal  denticles,  and  by  its  more  lunate 
tail.  It  also  grows  much  larger  than  the  Portu- 
guese shark.  We  need  only  note,  further,  that 
while  its  upper  teeth  are  narrow  and  awl-like,  its 
lowers  are  broad,  squarish,  forming  a  nearly 
continuous  cutting  edge,  with  the  single  cusp 
directed  sharply  outward;  that  its  gill  openings  are 
short  and  located  low  down  on  the  sides  of  the 
neck;  that  its  eyes  are  very  small;  and  that  it  is 
stout  shouldered,  with  blunt  rounded  snout,  as 
Scoresby  pictured  it  more  than  a  century  ago.43 

Color. — Blackish,  coffee  brown,  or  ashy-,  pur- 
plish-, or  slate  gray,  below  as  well  as  above; 
changing  to  bluish  gray  if  the  epidermis  is  rubbed 
off,  as  is  apt  to  happen  when  one  is  caught;  the 
back  and  sides  are  marked  with  many  indistinct 
dark  crossbars  on  some  specimens. 

Size. — This  is  one  of  the  larger  sharks.  It  is 
said  to  grow  to  a  length  of  24  feet,  but  21  feet  is  the 
largest  of  which  we  find  definite  record,44  and  16- 
to  18-footers  are  unusual.  One  of  16%  feet  was 
reported  from  the  Grand  Banks  in  1934;  one  of 

«  Arctic  Regions,  1820,  vol.  2,  pi.  15,  flgs.  3  and  4. 
"  Jenkins.  Fishes  British  Isles,  1925,  p.  325. 


16  feet  off  Portland,  Maine,  in  1846;  one  of  about 
15  feet  off  Cape  Ana  in  1849;  and  another  of  about 
that  same  size  was  caught  on  a  long  line  north  of 
Cape  Ann  in  February  1931.  Perhaps  8  to  14  feet 
is  a  fair  average  for  adults,  that  is  not  often  ex- 
ceeded among  the  hundreds  caught  annually  off 
West  Greenland  and  around  Iceland.  The  21- 
foot  British  specimen  mentioned  above  was  said 
to  weigh  about  2,250  pounds;  two  Gulf  of  Maine 
specimens,  each  about  1 1  feet  long,  weighed  about 
600  and  650  pounds,  respectively. 

Habits. — Off  Greenland,  and  along  the  Labrador 
coast,  the  Greenland  sharks  tend  to  approach  the 
surface  in  winter,  often  coming  right  up  to  the  ice. 
But  most  of  them  withdraw  in  summer  to  100 
fathoms  or  deeper.  And  the  few  that  visit  our 
Gulf  appear  to  hold  rather  closely  to  the  bottoms 
of  the  deeper  troughs,  though  a  stray  may  come 
so  close  to  the  shore  now  and  then,  and  intc  water 
so  shoal  as  to  blunder  into  a  fish  weir;  one  such 
event  is  on  record  for  Passamaquoddy  Bay. 

This  is  one  of  the  most  sluggish  of  sharks, 
offering  no  resistance  whatever  when  hooked,  and 
it  is  entirely  inoffensive  to  man.46     But  it  is  ex- 

"  Tales  to  the  effect  that  it  attacks  Qreenlanders  in  their  kyaks  are  appar- 
ently mythical,  and  Doctor  Porsild,  Director  of  the  biological  station  at 
Disko,  said  that  the  Eskimos  do  not  tear  it  as  they  do  the  killer  whale;  nor 
Is  there  any  authentic  instance  on  record  of  a  shark  attacking  a  human  being 
near  Iceland. 


FISHES   OF  THE  GULF  OF   MAINE 


55 


tremely  rapacious.  It  devours  any  carrion  ea- 
gerly, such  as  whale  meat,  blubber  from  whaling 
operations,  or  the  carcasses  of  young  seals  that 
are  left  on  the  ice  off  the  Newfoundland-Labrador 
coasts.  And  its  habit  of  gathering  when  there 
has  been  a  big  killing  of  narwhals  in  Greenland 
waters  is  proverbial.  Apart  from  carrion  (which 
cannot  be  available  except  on  rare  occasions),  its 
diet  includes  a  wide  variety  of  fishes,  large  and 
small.  Seals  are  a  favorite  food,  and  in  view  of 
its  sluggishness,  it  is  somewhat  astonishing  that 
it  should  be  able  to  capture  prey  as  active  as  seals, 
halibut,  and  salmon.  The  specimen  from  Cape 
Cod  Bay,  mentioned  above,  contained  half  a  dozen 
flounders  and  a  large  piece  (with  hide  and  hair) 
that  had  been  bitten  out  of  the  side  of  a  seal.  It 
is  also  known  to  eat  crabs,  large  snails,  even 
medusae.  Objects  as  large  as  an  entire  reindeer 
(without  horns),  a  whole  seal,  a  3-foot  cod,  and  a 
39-inch  salmon,  found  in  Greenland  shark 
stomachs,  give  some  measure  of  their  appetite. 
In  line  with  this,  they  will  bite  on  any  fish  or  meat 
bait,  the  more  putrid  and  ill  smelling  the  better. 

Large  numbers  of  soft  eggs,  without  horny  cap- 
sules, ranging  in  size  up  to  that  of  a  goose  egg, 
have  been  found  repeatedly  in  female  Greenland 
sharks,  but  never  any  embryos,  suggesting  that 
this  may  be  an  egg-laying  species.46 

General  range. — Northern  Atlantic,  from  Polar 
latitudes  south  to  the  North  Sea  and  accidentally 
to  the  mouth  of  the  Seine  and  perhaps  to  Portugal 
in  the  east;  south  regularly  to  Newfoundland  and 
the  northern  part  of  the  Gulf  of  St.  Lawrence  in 
the  west,  and  less  commonly  to  the  Gulf  of  Maine. 
It  is  represented  in  the  Mediterranean  region,  in 
the  North  Pacific,  and  in  the  sub-Antarctic  by 
forms  that  appear  to  be  distinct,  though  closely 
allied  to  it.47 

Occurrence  in  the  Gulf  of  Maine. — Although  there 
is  no  reason  to  suppose  that  the  Greenland  shark 
ever  appears  in  our  Gulf  save  as  a  straggler  from 
the  north,  its  presence  there  has  been  signalized 
on  a  number  of  occasions.  Two  specimens,  for 
example,  were  taken  in  the  neighborhood  of  St. 
Andrews  in  1915  (one  caught  in  a  weir  and  the 
other  on  a  long  line).  It  has  been  reported  off 
Eastport ;  off  Cape  Elizabeth  whence  6  were  landed 


*•  The  Mediterranean  Somniosut  Tostratus,  on  the  contrary,  bears  living 
young. 

*7  For  recent  discussion  of  the  species  of  Somniosus,  see  Bigelow  and 
Schroeder.  Fishes  Western  North  Atlantic,  Pt.  1,  1948,  p.  515. 


at  Portland  between  1925  and  1948;48  on  Jeffreys 
Ledge,  where  one  of  about  15  feet  was  caught  on 
a  long  line,  on  February  16,  1931  ;40  near  Cape 
Ann;  off  Marblehead  and  Nahant;  in  Massachu- 
setts Bay;  off  Barnstable  in  Cape  Cod  Bay;  at 
Provincetown ;  and  in  Cape  Cod  Bay  off  the  en- 
trance to  the  Cape  Cod  Canal,  where  one  between 
10  and  11  feet  long  was  taken  by  a  trawler  in 
April  1924,  landed  in  Boston  and  identified  by  us. 

Recorded  captures  in  the  Gulf  include  small 
specimens  as  well  as  large,  and  have  been  for  all 
four  seasons  of  the  year,  suggesting  that  when  a 
Greenland  shark  does  stray  southward  to  the 
Gulf,  it  may  survive  there  for  years.  The  local 
records  are  distributed  so  widely  as  to  show  that 
an  odd  specimen  is  to  be  expected  anywhere  in 
the  deeper  parts  of  the  Gulf.  And  rumor  has  it 
that  they  were  more  numerous  in  our  waters  in 
early  colonial  times  when  Atlantic  right  whales 
were  still  being  killed  in  numbers  off  the  Massa- 
chusetts coast.60 

Commercial  importance.- — This  shark  is  not  plen- 
tiful enough  in  our  Gulf  to  be  even  of  potential 
value.  But  it  has  long  supported  a  fishery  off 
northern  Norway,  around  Iceland,  and  in  West 
Greenland  waters,  chiefly  for  its  liver  oil.61  In 
Greenland  the  flesh  is  dried  also  for  dog  food,  and 
to  a  small  extent  in  Iceland  for  human  consump- 
tion. But  it  produces  an  intoxicant  poisoning  if 
eaten  fresh,  though  it  is  wholesome  if  dried.62 

Dalatias  licha  (Bonnaterre)  1788 

Bigelow  and  Schroeder,  1948,  p.  502. 

Description. — This  shark  resembles  the  Portu- 
guese shark  in  the  relative  sizes  and  positions  of 
its  fins;  also  in  its  scales.  But  its  dorsal  fins  do 
not  have  any  trace  of  spines,  while  the  serrate 
margins  of  its  lower  teeth,  in  combination  with 
their  triangular  shape,  mark  it  off  from  any  other 
shark  without  an  anal  fin  that  is  known  yet  from 
the  North  Atlantic.  Its  trunk  is  rather  slender, 
its  snout  short  and  bluntly  rounded,  and  the 
lower-anterior  corner  of  its  tail  fin  is  not  expanded 
as  a  definite  lobe.     Its  upper  teeth  are  slender,  awl- 

"  Reported  to  us  by  the  late  W.  W.  Rich. 

19  This  one  was  landed  in  Boston,  where  we  saw  it. 

»  When  they  gather  to  feed  on  whale,  narwhal,  and  seal  carcasses  in  their 
northern  home,  they  may  linger  for  a  long  time  in  the  vicinity. 

"  The  annual  catch  off  West  Greenland  was  around  32,000  during  the  first 
decade  of  the  present  century. 

M  For  accounts,  see  Jensen,  1914  (Selachians  of  Greenland,  Mlndesk. 
Jap.  Steenstrup,  vol.  2,  No.  30,  1914,  p.  12);  also  Clark  (Science,  N.  Ser., 
vol.41, 1915,  p.  795). 


56 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


i\/\/\/\AA 


Figure  22. — Dalatias  (Dalatias  licha),  female,  58  inches  long,  from  Georges  Bank.    A,  upper  teeth  and  B,  lower  teeth 
from  central  part  of  mouth,  about  1.5  times  natural  size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


shaped,  curving  somewhat  outward  toward  the 
corners  of  its  mouth;  but  the  lowers  are  erect, 
broadly  triangular,  with  serrate  edges. 

Color. — Dark  chocolate,  cinnamon,  or  violet 
brown  below  as  well  as  above;  the  upper  surface 
sometimes  with  poorly  defined  blackish  spots;  the 
dorsal  and  pectoral  fins  with  pale  or  whitish  edges, 
the  tail  tipped  with  black. 

Size. — Most  of  those  caught  are  between  40 
and  60  inches  long;  72  inches  is  the  longest  re- 


corded so  far.  The  Gulf  of  Maine  specimen  illus- 
trated in  figure  22  was  about  5  feet  long  and 
weighed  23%  pounds,  gutted. 

General  range. — Eastern  Atlantic,  from  tropical 
West  Africa  to  the  Irish  Atlantic  slope;  recorded 
once  from  the  American  coast. 

Occurrence  in  the  Gulf  of  Maine. — Our  only  rea- 
son for  mentioning  this  shark  is  that  a  female, 
about  5  feet  long,  was  taken  on  the  northern  edge 
of  Georges  Bank  on  August  19,  1937  (fig.  22)  ,63 


THE  BRAMBLE  SHARKS.     FAMILY  ECHINORHINIDAE 


The  only  living  representative  of  this  family  (it 
is  represented  among  the  tertiary  sharks)  re- 
sembles the  Greenland  shark  and  its  allies  in 
lacking  both  anal  fin  and  dorsal  spines,  but  its 
teeth  are  alike  in  the  two  jaws. 

Bramble  shark  Echinorhinus  brucus 
(Bonnaterre)  1788 

Spiny  shark 

Bigelow  and  Schroeder,  1948,  p.  527. 


Description. — The  location  of  the  first  dorsal  fin 
above  the  pelvics  instead  of  about  midway  between 
the  latter  and  the  pectorals,  and  the  very  different 
shape  of  its  tail  fin  (cf.  fig.  23  with  fig.  21),  are  the 
most  conspicuous  field  marks  separating  this  shark 
from  the  Greenland  shark.  Brucus  also  differs 
from  the  latter  in  that  the  teeth  are  alike  in  the 
two  jaws,  instead  of  unlike,  and  that  the  skin  of  its 
back  and  sides  is  sparsely  strewn  with  large  scales 
with  either  one  or  two  sharp  points. 

«  Recorded  by  Nichols  and  Firth,  Proc.  Biol.  Biol.  Soc.  Wash.,  vol.  62, 
1939,  p.  85. 


•■  Q  9  w  « 


Figure  23. — Spiny  shark  (Echinorhinus  brucus),  eastern  Atlantic  specimen  about  3  feet  long. 

der.     Drawing  by  W.  P.  C.  Tenison. 


From  Bigelow  and  Schroe- 


FISHES    OF  THE   GULF   OF    MAINE 


57 


Color. — Described  as  dark  gray,  olive  or  brown 
above,  with  metallic  reflections,  and  with  or 
without  darker  blotches;  as  paler  brown  or  gray 
to  white  below.  The  scales  have  been  described 
as  luminescent,64  but  there  are  no  special  luminous 
organs. 

Size. — The  largest  of  which  we  have  found  a 
record  (a  specimen  from  British  waters)  was  9 
feet  long.  One  8  feet  4  inches  long  weighed  about 
300  pounds. 

General  range.- — Eastern  Atlantic  (including  the 
Mediterranean)    from    tropical    West    Africa    to 


Ireland  and  the  North  Sea,  and  accidental  in  the 
western  Atlantic;  represented  in  South  Africa; 
off  California;  in  the  Hawaiian,  Japanese,  and 
Australo-New  Zealand  regions,  and  in  Arabian 
waters  by  forms  that  probably  cannot  be  dis- 
tinguished from  brucus  of  the  Atlantic. 

Occurrence  in  the  Gulf  of  Maine. — A  single 
specimen  of  this  little  known  shark  came  ashore 
at  Provincetown  in  December  1878.  This  and 
one  taken  near  Buenos  Aires  more  recently  M 
are  the  only  records  of  it  from  the  western  Atlantic. 


Torpedoes,  Skates,  and  Rays.     Order  Batoidei 


This  tribe  falls  into  four  groups,  so  far  as  the 
Gulf  of  Maine  fauna  is  concerned:  first,  the 
torpedoes  (family  Torpedinidae) ,  with  large  caudal 
fin,  interesting  because  provided  with  electric 
organs  capable  of  giving  a  strong  shock;  second, 
the  skates  (family  Rajidae),  with  very  thin  bodies, 
comparatively  short  tails  without  tail  spines,  and 
only  a  trace  of  caudal  fin;  third,  the  sting  rays 
(families  Dasyatidae  and  Rhinopteridae) ,  with 
long  whiplike  tails  armed  with' a  stiff  saw-edged 
spine    (or  spines);   and   fourth,    the    devil   rays 

«  Cornish,  Zoologist,  Ser.  2,  vol.  10, 1875,  p.  4801. 


(Mobulidae)  with  two  ear-like  fins  extending 
forward  from  the  front  of  the  head.  Most  of  our 
common  species  belong  to  the  second  group. 

Among  torpedoes,  skates,  and  rays,  fertiliza- 
tion is  internal  as  it  is  among  sharks,  and  the 
modification  of  the  posterior  edges  of  the  pelvic 
fins  into  rodlike  semitubular  claspers  (the  copula- 
tory  organs)  distinguishes  males  and  females  at  a 
glance.  Some  bear  "living"  young,  ready  for 
independent  existence;  others  lay  eggs. 

"  Berg,  Com.  Ictiol.  Comm.  Mas.  Nac.  Buenos  Aires,  vol.  1,  No.  1,  1898, 
p.  10. 


KEY  TO  GULF  OF  MAINE  SKATES  AND  RAYS 

1.  The  front  of  the  head  bears  a  pair  of  separate,  ear-like  fins,  extending  forward Devil  ray,  p.  77 

The  front  of  head  does  not  bear  a  pair  of  separate  ear-like  fins  extending  forward 2 

2.  There  is  a  large  triangular  caudal  fin  as  well  as  two  well  developed  dorsal  fins  on  the  tail Torpedo,  p.  58 

There  is  no  distinct  caudal  Ad;  the  dorsal  fins,  if  any,  are  very  small 3 

3.  No  long  dorsal  spine  on  tail Common  skates  4 

There  is  a  long  saw-edged  dorsal  spine  (or  spines)  on  the  tail 11 

4.  The  upper  surface  of  the  disc  is  marked  with  conspicuous  black  rosettes Leopard  skate,  p.  66 

The  markings  on  the  upper  surface  of  the  disc  are  not  in  the  form  of  black  rosettes 5 

5.  There  are  no  conspicuous  thorns  along  the  mid-dorsal  zone  of  disc  between  the  spiracles  and  the  base  of  tail;  the  lower 

surface  of  disc  is  marked  with  black  dots  or  dashes,  marking  the  openings  of  the  mucous  pores. 

Barndoor  skate,  medium  sized  and  large  specimens,  p.  61 

There  are  one  or  more  rows  of  conspicuous  thorns  along  the  mid-dorsal  zone  of  disc  rearward  from  the  spiracles;  the 

lower  surface  of  disc  is  not  marked  with  black  dots  or  dashes 6 

6.  There  are  no  large  thorns  on  the  rear  Y*-Yi  of  tail Smooth-tailed  or  Prickly  skate,  p.  70 

There  are  one  or  more  rows  of  large  thorns  along  the  rear  part  of  tail  as  well  as  farther  forward  along  it 7 

7.  There  are  no  large  thorns  on  upper  side  of  disc  between  the  spiracles  and  the  level  of  axils  of  pectoral  fins. 

Barndoor  skate,  very  small  specimens,  p.  61 
The  upper  side  of  disc,  rearward  from  spiracles,  has  more  or  fewer  large  thorns 8 

8.  The  thorns  of  the  midrow  on  the  tail  are  much  larger  and  more  conspicuous  than  any  other  thorns  on  the  tail,  and  not 

more  than  9  or  10  in  number Thorny  skate,  p.  72 

No  one  row  of  thorns  along  the  tail  is  much  larger  or  more  conspicuous  than  the  other  thorns  on  the  tail ;  there  are  at 

least  15  thorns  in  each  of  the  rows  along  tail 9 

210941 — 53 5 


58 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


9.  There  is  only  one  row  of  large  thorns  along  the  midzone  of  the  disc  from  the  nape  to  the  level  of  the  axils  of  the  pectoral 

fins;  the  first  and  second  dorsal  fins  are  separated  by  a  definite  space  or  at  least  by  1  or  2  thorns;  the  forward  angle 
of  the  disc  is  less  than  110°;  the  upper  surface  of  the  disc  is  marked  with  short  dark  bars  as  well  as  with  roundish 

spots Brier  skate,  p.  65 

There  are  at  least  three  rows  of  thorns  along  the  midzone  of  the  disc  from  the  nape  to  the  level  of  the  axils  of  the 
pectoral  fins;  the  first  and  second  dorsal  fins  are  not  separated  by  a  definite  interspace  or  by  a  thorn  or  thorns;  the 
forward  angle  of  the  disc  is  more  than  125°;  the  upper  surface  is  not  marked  with  dark  bars  though  it  is  variously 
spotted 10 

10.  Upper  teeth  in  at  least  72  series,  most  often  90-100;  does  not  mature  sexually  until  at  least  26  inches  long. 

Big  skate,  p.  63 
Upper  teeth  in  not  more  than  66  series  and  usually  less  then  54;  matures  when  only  18-20  inches  long. 

Little  skate,  p.  67 

11.  There  is  a  small  dorsal  fin  on  the  upper  side  of  the  tail,  in  front  of  the  spine  (or  spines) ;  the  crown  of  the  head  is  high- 

domed,  with  the  eyes  and  spiracles  on  the  sides;  there  are  only  7-9  series  of  teeth  in  the  form  of  large  flat  grinding 

plates - Cow  nosed  ray,  p.  76 

There  is  no  dorsal  fin  on  the  tail;  the  crown  is  low,  flat,  and  with  the  eyes  and  spiracles  on  the  upper  surface;  the 
teeth  are  in  many  series,  in  mosaic  arrangement Sting  ray,  p.  74 

THE  TORPEDOES  OR  ELECTRIC  RAYS.  FAMILY  TORPEDINIDAE 


The  trunk  of  the  electric  rays  has  the  form  of  a 
flattened,  roundish  or  oval  disc,  fleshier  toward 
the  margins  than  it  is  in  other  Gulf  of  Maine 
skates  or  rays,  and  the  body  is  softer.  The  tail, 
too,  is  broader  and  shorter;  there  are  one  or  two 
relatively  larger  dorsal  fins  on  the  tail,  and  the 
latter  ends  in  a  well-developed  caudal  fin  also. 

The  most  interesting  feature  of  the  electric  rays 
is  that  they  have  two  large  electric  organs,  each 
of  which  occupies  one  side  of  the  front  part  of  the 


disc.  In  the  only  Gulf  of  Maine  species  the  two 
organs  together  make  up  about  one-sixth  of  the 
total  weight  of  the  fish. 

Torpedo  Torpedo  nobiliana  Bonaparte  1835 

Electric  ray;  Numbfish,  Crampfish 

Bigelow  and  Schroeder,  1953,  p.  96. 

Garman,  1913,  pi.  25,  fig.  2,  as  Narcacion  nobilianus. 

Description. — No  one  would  be  apt  to  mistake  a 


Figttbe  24. — Torpedo  (Torpedo  nobiliana),  male,  about  33  inches  long,  off  Plymouth,  Massachusetts.     A,  side  view  of 
caudal  fin;  B,  teeth  3  times  natural  size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


FISHES   OF  THE  GULF   OF   MAINE 


59 


torpedo  for  any  other  Gulf  of  Maine  skate  or  ray, 
the  rounded  outline  of  the  disk  and  the  large 
caudal  fin  identifies  it  at  a  glance.  Furthermore, 
its  skin  is  soft  and  naked,  without  the  spines  or 
thorns  so  characteristic  of  all  our  common  skates. 
The  disk  is  roughly  subcircular,  truncate  in  front, 
and  somewhat  broader  than  long.  The  eyes  are 
very  small  and  set  far  forward.  The  two  dorsal 
fins,  of  which  the  first  is  the  larger,  stand  on  the 
forward  end  of  the  tail,  the  first,  indeed,  partly 
above  the  bases  of  the  pelvic  fins,  and  they  are 
separated  by  an  interspace  nearly  as  long  as  the 
second  dorsal  fin.  The  tail  fin  is  of  ordinary  fish 
form,  triangular  and  nearly  as  long  as  it  is  deep. 
The  tail  is  shorter  than  in  the  skates  for  it  occupies 
only  about  two-fifths  the  total  length  of  the  fish, 
measured  from  the  cloaca.  The  teeth  are  small, 
with  sharp  curved  points,  and  are  in  about  60 
series,  with  up  to  7  rows  exposed  and  functioning 
at  one  time. 

Color. — Dark  chocolate  to  purplish  brown 
above,  some  with  a  few  obscure  darker  spots; 
lower  surface  white  except  that  the  edges  of  disk, 
fins,  and  tail  are  of  the  same  dark  tint  as  the 
upper  side. 

Size. — Adult  torpedoes  are  usually  2  to  5  feet 
long  or  a  little  longer,  and  heavy  for  their  size. 
Specimens  taken  at  Woods  Hole  average  about  30 
pounds,  while  most  of  those  taken  anywhere  on 
our  Atlantic  coast  weigh  less  than  75  pounds. 
But  we  have  seen  one  only  about  4  feet  long  from 
Chesapeake  Bay  that  weighed  about  100  pounds; 
one  of  144  pounds  was  brought  from  Nantucket 
to  the  U.  S.  Fisheries  Station  at  Woods  Hole 
many  years  ago;  and  the  heaviest  taken  near 
Provincetown  were  estimated  long  ago  by  a 
fisherman  of  keen  observation  as  170  to  200 
pounds. 

Habits. — The  most  interesting  thing  about  the 
torpedo  is  its  ability  to  give  electric  shocks  of 
considerable  strength  to  anyone  touching  it. 
The  statement,  even,  has  long  been  current  that 
the  shock  from  a  large  one  in  rested  condition  may 
be  strong  enough  to  throw  a  full  grown  man  to  the 
ground.  And  the  story  is  told  of  a  dog  which  was 
in  the  habit  of  wading  on  a  Cape  Cod  beach  in 
shoal  water  to  catch  flounders,  but  was  so  shocked 
by  a  torpedo  that  it  ran  away  howling  and  could 
never  be  persuaded  to  go  fishing  again.  In  fact, 
this  anecdote  antedates  the  scientific  naming  of 
the   New   England    torpedo.     But   shocks   of   a 


strength  even  approaching  what  is  suggested  by 
such  reports  are  to  be  expected  only  from  torpedos 
of  the  largest  size  in  rested  condition.  The  voltage 
recorded  recently  was  170  to  220  for  one  that  had 
been  kept  in  a  live  well.  And  the  most  we  have 
felt  ourselves  from  medium-sized  torpedos  lying 
on  the  dock  at  Woods  Hole  has  been  a  slight 
benumbing  sensation. 

The  torpedo,  like  others  of  its  tribe,  is  a  bottom 
fish.  It  is  a  fish  eater.  The  stomach  of  one  taken 
at  Woods  Hole  contained  a  summer  flounder 
(Paralichthys  dentatus)  about  14K  inches  long.  A 
2-pound  eel,  a  1-pound  flounder,  plaice  (Pleuro- 
nectes  platessa),  red  mullet  {Mullus  surmuletus),  a 
salmon  weighing  4  or  5  pounds,  and  the  remains 
of  spotted  dogfish  (genus  Scyliorhinus)  have  been 
found  in  the  stomachs  of  British  specimens.  The 
wide  distensibility  of  its  jaws  allows  it  to  swallow 
fishes  much  larger  than  might  be  considered 
possible  from  the  breadth  of  the  mouth  when 
closed.  And  it  is  generally  believed  that  it  stuns 
its  prey  by  its  electric  shocks.  Otherwise  it  is 
difficult  to  conceive  how  so  sluggish  a  fish  could 
capture  such  active  prey. 

It  bears  "living"  young,  but  there  is  no  placen- 
tal connection  between  embryo  and  mother. 
And  it  seems  that  the  young  are  born  offshore,  for 
the  smallest  torpedo  yet  recorded  from  American 
inshore  waters  (from  New  Jersey)  was  about  2 
feet  (610  mm.)  long.  And  we  doubt  if  it  succeeds 
in  producing  young  in  the  colder  waters  of  our 
Gulf. 

General  Range. — Both  sides  of  the  North  Atlan- 
tic M  from  southern  Nova  Scotia  (La  Have  Bank), 
Bay  of  Fundy,  and  Georges  Bank  to  North 
Carolina  in  the  west  ;w  and  from  northern  Scotland 
to  the  Mediterranean,  Azores,  Madeira,  and 
tropical  West  Africa  in  the  east. 

Occurrence  in  the  Gulf  of  Maine. — -The  torpedo  is 
more  common  south  and  west  from  Cape  Cod 
than  to  the  northward  and  eastward.  But  it 
strays  past  the  elbow  of  the  Cape  often  enough 
for  it  to  be  classed  as  a  regular  member  of  the 
Gulf  of  Maine  fish  fauna.  The  most  northeasterly 
records  for  it  are  of  one  presumably  of  this  species 
taken  in  St.  Margarets  Bay,  Nova  Scotia,  some  30 
years  ago;  one  caught  on  a  long  line  set  for  cod 


"  Comparison  of  American  specimens  with  one  from  the  North  Sea  revealed 
no  differences. 

•'  This  torpedo  is  also  reported  from  the  Florida  Keys  and  from  Cuba,  but 
on  doubtful  evidence. 


60 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE    SERVICE 


on  La  Have  Bank  in  1890,68  and  from  Eastport, 
Maine,  at  the  mouth  of  the  Bay  of  Fundy.  It 
has  also  been  taken  at  Williamsport,  Maine;  off 
Seguin  Island  where  one  was  examined  in  1880; 
at  the  mouth  of  Casco  Bay;  at  Wood  Island  near 
Cape  Elizabeth  (1,  in  a  trap,  in  1894);  near  Cape 
Ann;  off  Plymouth  in  the  southern  side  of  Massa- 
chusetts Bay;  near  Provincetown;  and  on  the 
outer  coast  of  Cape  Cod,  so  it  would  be  no  sur- 
prise to  find  it  anywhere  along  the  shores  of  the 
Gulf.  It  has  been  caught  occasionally  on  Georges 
Bank;69  there  are  records  of  long  standing  of 
torpedos  off  Nantucket  and  Marthas  Vineyard, 
and  they  are  caught  yearly  in  Vineyard  Sound 
and  in  Buzzards  Bay. 

Most  of  the  reports  of  torpedoes  within  the 
Gulf  have  been  based  on  single  specimens.  But  it 
has  been  known  for  a  long  time  that  torpedoes 
are  caught  in  much  larger  numbers  in  some  years 
than  in  others.  Thus  they  are  said  to  have  been 
unusually  common  near  Provincetown  in  1819 
and  for  the  next  4  or  5  years,  when  60  to  80  were 
taken  there  yearly.     Again  in  1845  about  a  dozen 


came  ashore  or  were  caught  otherwise  near 
Provincetown.  Any  fluctuation,  however,  that 
may  have  taken  place  from  year  to  year  thereafter 
seems  to  have  attracted  no  attention  until  the 
summer  of  1896,  when  Dr.  W.  C.  Kendall,  of  the 
U.  S.  Fish  Commission  collected  several  along  the 
coast  of  Maine.  The  Massachusetts  Bay  speci- 
men mentioned  above,  taken  off  Plymouth  and 
now  in  the  Harvard  Museum  of  Comparative 
Zoology,  is  the  only  torpedo  from  the  inner  part 
of  the  Gulf  of  which  we  have  heard  since  that 
time.  But  it  is  as  likely  to  be  found  in  the  Gulf 
now  as  it  ever  was. 

Importance. — The  torpedo  is  of  no  commercial 
value  nowadays,  but  its  liver  oil  was  considered 
equal  to  the  best  sperm  for  illuminating  purposes 
before  the  use  of  kerosene  oil  was  general.  There 
is  an  old  tale  that  its  oil  was  a  good  cure  for 
cramps  if  rubbed  on  externally,  for  stomach 
trouble  if  taken  internally.  And  when  one  is 
landed  on  the  dock  at  Woods  Hole  it  is  an  object 
of  interest  to  the  workers  at  the  Biological  Labora- 
tory because  of  its  electric  discharges. 


SKATES.     FAMILY  RAJIDAE 


Skates,  with  their  disc-like  outlines,  thin  as  a 
shingle  toward  their  outer  edges,  and  with  their 
rather  long  tails,  are  familiar  objects  along  our 
shores.  The  outer  edges  of  their  pelvic  fins  are 
concave  (convex  in  the  sting  rays) ,  they  have  two 
very  small  dorsal  fins  on  the  rear  part  of  the  tail, 
but  no  distinct  tail  fin,  and  they  lack  the  large 
tail  spine  that  is  so  characteristic  of  the  sting  rays. 
The  Gulf  of  Maine  supports  four  species  in  abun- 
dance, while  two  others  have  been  recorded  on 
rare  occasions. 

The  common  skates  look  so  much  alike  that 
fishermen  seldom  distinguish  between  them.  For 
this  reason  we  know  very  little  about  the  indi- 
vidual differences  in  habits  among  the  several 
species.  All  live  chiefly  on  the  bottom  or  close  to 
it,  spending  much  of  the  time  partially  buried  in 
the  mud  or  sand.  They  move  through  the  water 
by  undulations  of  the  flexible  pectoral  fins,  steering 
themselves  with  the  tail.  All  are  decidedly  om- 
nivorous, feeding  largely  on  the  larger  Crustacea, 

"  Reported  by  Q.  F.  O.  Hansen,  then  second  mate  and  later  master  of  the 
U.  S.  Fish  Commission  schooner  Oramput,  who  doubtless  was  acquainted 
with  the  torpedo  at  Woods  Hole. 

•>  The  most  recent  record  is  of  one  58  inches  long,  trawled  on  the  southwest 
part  in  December  1930. 


such  as  shrimps,  crabs,  lobsters;  on  mollusks, 
worms,  and  to  a  greater  or  less  extent  on  fish. 

All  the  true  skates  lay  large  eggs  with  blackish 
or  sea-green  leathery  shells,  roughly  oblong  in 
outline,  with  a  hollow  tendril  at  each  corner  by 
which  they  become  fastened  to  seaweeds  or  other 
objects.  The  empty  eggshells, " mermaids  purses," 
are  familiar  objects  on  our  beaches  among  the 
flotsam  along  high  water  mark.  While  still  in 
the  egg  the  embryo  skate  develops  temporary 
external  gill  filaments  from  the  walls  of  the  gill 
clefts,  but  these  disappear  completely  before  it 
hatches.  Probably  all  our  local  skates  spawn 
over  a  considerable  part  of  the  year,  with  incuba- 
tion periods  of  several  months  up  to  a  year  or 
more.60 

To  give  some  idea  of  their  abundance  on  the 
offshore  banks  we  may  note  that  the  average 
number  of  skates  (all  species  together)  taken  on 
Georges  Bank,  per  trip  of  4  to  7  days,  on  25 
trips  by  several  trawlers,  January  to  December 
1913,  was  about  800,  the  largest  catch  4,520,  the 

w  Under  aquarium  conditions  the  incubation  period  for  the  little  skate 
(R.  trinacea)  was  5  to  6  months  (p.  69);  and  it  ranged  from  4H  to  about  HH 
months  for  6  common  European  skates;  see  Clarke,  Jour.  Marine  Biol. 
Assoc.  United  Kingdom,  vol.  12,  No.  4,  1927,  p.  687. 


FISHES   OF  THE  GULF  OF   MAINE 


61 


poorest  82.  Again,  on  a  trip  to  the  northeastern 
part  of  the  bank,  September  1929,  on  the  otter 
trawler  Kingfisher,  37  hauls  yielded  from  0  to  105 
skates  per  haul  (total  495)  and  42  trawl  hauls  by 
the  Eugene  H,  fishing  from  Nantucket  Lightship 
to  the  south-central  part  of  Georges  Bank  in  late 
June  1951  caught  an  average  of  146  skates  per 
haul  (total,  6,130  skates),  which  works  out  at 
about  9  to  10  skates  per  acre.61  Probably  they 
are  equally  abundant  on  Browns  Bank;  certainly 
they  are  familiar  enough  there,  but  statistics  are 
not  available  of  the  actual  numbers  caught. 
Skates  are  also  plentiful  inshore  as  appears  from 
catches  of  about  1  skate  to  33  fishes  of  all  kinds 
on  long  lines,  at  various  localities  in  the  Gulf  of 
Maine.62 

In  the  Gulf  of  Maine,  skates  are  only  a  nuisance 
for  they  bite  the  hook  readily  and  often  are  caught 
in  great  numbers  in  otter  trawls,  most  of  them  to 
be  thrown  back  into  the  sea,  the  market  demand 
for  them  being  so  small  that  the  total  landings 
reported  for  New  England  (Massachusetts  and 
Maine)  in  1947  was  only  28,200  pounds;  and 
59,100  pounds  for  1948.  But  some  are  now  being 
landed  in  Maine  for  fish  meal.63  They  are  much 
more  highly  valued  in  northwesternEurope  for  food 
with  landings  for  the  years  just  preceding  World 
War  II,  running  around  90  to  100  million  pounds. 

Barn-door  skate  Raja  laevis  Mitchill  1817 

Bigelow  and  Schroeder,  1953,  p.  217. 
Garman,  1913,  pi.  22,  fig.  2,  as  R.  stabuliforis. 

Description.- — The  barn-door  skate  is  easily 
identified  by  its  large  size,  its  very  pointed  snout, 
and  its  smooth  skin.  The  thorns  along  the  mid- 
line of  its  back  are  comparatively  small  and  run 
only  from  the  hinder  part  of  the  disc  back  along 
the  tail ;  the  tail  also  has  one  or  two  rows  of  large, 
sharp  spines  (smaller  on  males  than  on  females) 
along  each  side,  besides  the  median  row.  There 
are  small  thorns  on  the  snout  also,  sometimes 
below  as  well  as  above,  and  along  the  front  edges 
of  the  pectoral  fins.  The  male  has  a  patch  of 
erectile  hooks  on  the  outer  part  of  each  pectoral 
covering  an  area  measuring  5  by  1%  inches  on  one 
side,  and  4%  by  1%  inches  on  the  other  in  a  speci- 

11  Three  mile  hauls  with  the  trawl  sweeping  a  strip  about  35  feet  wide. 

•'  Examples  are:  15  miles  off  Monhegan  I.,  Maine,  June  24-25,  1913,  total 
fish  caught,  5,463;  skates  170.  Twenty  miles  east  of  Cape  Cod,  Nov.  11, 1913; 
total  fish  caught  6,532,  skates  202.  Jeffreys  Ledge,  Dec.  11-12,  1913;  total 
fish  caught  3,996,  skates  62. 

B  Scattergood,  Copeia,  1950,  p.  169. 


men  52  inches  long;  otherwise  the  pectorals  are 
smooth  for  the  most  part.  The  front  angle  of  the 
disc  is  sharper  than  in  our  other  skates,  being 
more  acute  than  a  right  angle,  but  the  tip  of  the 
snout  is  blunt.  The  outer  corners  of  the  pectorals 
are  angular  and  the  disc  as  a  whole  is  diamond  or 
lozenge-shaped.  The  two  dorsal  fins  are  separated 
by  a  short  interspace,  with  one  or  more  spines, 
and  the  tip  of  the  tail  extends  farther  beyond  the 
second  dorsal  fin  than  it  does  in  most  skates. 
The  teeth  of  the  female  are  flat  and  pavement- 
like, but  those  of  adult  males  are  provided  with 
sharp  slender  cusps.  Thirty  to  forty  series  of 
teeth  have  been  counted  in  the  upper  jaw,  28  to 
38  series  in  the  lower  jaw. 

Color. — The  barn-door  skate  like  so  many  sea 
fish,  varies  in  color.  The  upper  surface  is  brown 
(as  a  rule  usually  of  a  distinctly  reddish  hue), 
variously  marked  with  small  scattered  darker 
spots  or  blotches  of  varying  size,  and  often  with 
pale  marblings  or  waterings;  usually  there  is  a 
large  oval  spot  on  the  base  of  each  pectoral  fin,  in 
line  with  the  outer  angle.  The  lower  surface  is 
not  as  uniformly  pale  as  it  is  in  most  skates, 
its  gray  or  white  ground  being  shaded  with  darker 
toward  the  snout,  and  speckled  on  one-third  grown 
specimens  and  larger,  with  black  or  dusky  dots 
or  short  streaks  that  mark  the  mucous  pores, 
a  conspicuous  feature. 

Size. — The  barn-door  skate  is  our  largest, 
growing  to  a  length  of  5  feet;  it  has  been  said  to 
reach  6  feet  though  there  is  no  definite  record  of 
one  that  large.  One  of  58  inches  was  42  inches 
wide  with  a  tail  27  inches  long,  and  a  female  of 
50  inches,  taken  by  us,  was  33%  inches  wide,  with 
a  22-inch  tail.  Barn-door  skates  weigh  about  4 
to  6  pounds  when  28  to  30  inches  long,  about  10 
to  11  pounds  at  36  inches,  and  about  19  to  21 
pounds  at  45  to  46  inches.  Very  small  specimens 
are  seldom  taken. 

Habits.- — Barn-door  skates  are  bottom  fish. 
They  prefer  smooth  to  rocky  ground,  and  we 
have  caught  them  on  very  soft  mud  bottoms  as 
well  as  on  sand  and  gravel.  The  fact  that  the 
lower  surface  is  more  or  less  pigmented  instead 
of  white  suggests  that  it  hugs  the  bottom  less 
closely  than  other  skates,  and  it  is  a  strong, 
active  swimmer,  as  anyone  will  agree,  who  has 
landed  a  large  one  on  a  hand-line.  Its  usual  depth 
range  is  from  close  to  the  tide  line,  down  to  about 
100  fathoms.     It  is  perhaps  more  plentiful  at  25 


62 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  25. — Barn-door  skate  (Raja  laevis).  A,  dorsal  view  of  female,  about  47  inches  long,  Massachusetts;  B,  ventral 
view  of  one  of  about  26%  inches  to  show  the  black  markings;  C,  upper  teeth  from  center  of  jaw  of  female  50  inches  long; 
and  D,  upper  teeth  from  center  of  jaw  of  male  52  inches  long.  B,  C,  and  D  from  Nantucket  Shoals.  From  Bigelow  and 
Schroeder.     Drawings  by  E.  N.  Fischer. 


to  35  fathoms  on  Georges  Bank  and  on  Nantucket 
Shoals  than  deeper,  judging  from  average  catches 
of  32  per  haul  at  26  to  35  fathoms,  13  per  haul  at 
36  to  49  fathoms,  and  6  per  haul  at  50  to  75 
fathoms  in  42  trawl  hauls  by  the  Eugene  H,  late 
June  1951,  fishing  from  Nantucket  Lightship  to 
the  south-central  part  of  Georges  Bank.  But  the 
Atlantis  found  it  widespread  (though  not  numer- 
ous), as  deep  as  100  fathoms  both  in  the  open 
trough  of  our  gulf  and  in  the  bowl  west  of  Jeffreys 
Ledge  during  experimental  trawling,  in  August 
1936;  and  it  has  been  reported  as  deep  as  235 
fathoms  off  Nantucket. 

The  temperature  range  of  the  barn-door  skate 
is  wider  than  that  of  the  little  skate  (p.  67).  They 
are  found  in  the  southern  side  of  the  Guff  of  St. 
Lawrence  in  the  icy-cold-bottom  water  on  the 
banks,  also,  at  lesser  depths  that  warm  in  summer 
to  60°  F.  (16°  C.)  or  more.    In  the  Gulf  of  Maine, 


at  one  locality  or  depth  or  another,  they  are  ex- 
posed to  temperatures  ranging  from  perhaps  as 
low  as  32°  to  as  high  as  64  to  68°  and  the  upper 
limit  must  be  considerably  higher  in  the  southern 
part  of  their  range. 

Garman  has  pointed  out  that  the  spines  on  the 
snout  of  this  skate  are  usually  worn  smooth,  as 
though  used  to  dig  in  the  mud  or  sand  (very  likely 
it  thus  obtains  the  bivalves  that  form  part  of 
its  diet).  It  also  feeds  on  worms,  various  crus- 
taceans, particularly  on  large  rock  crabs  and  lob- 
sters, shrimps,  squid,  and  on  fish.  Probably  it  is 
more  destructive  to  the  latter  than  are  any  other 
of  our  skates  thanks  to  its  large  size.  Woods  Hole 
records  list  spiny  dogfish,  alewives,  herring,  men- 
haden, butterfish,  launce,  cunners,  tautog,  scul- 
pins,  silver  hake,  hake,  and  flatfish  among  its 
foods.  No  doubt  cod,  haddock,  and  other  fish, 
suffer  to  some  extent  from  this  skate  on  the  off- 


FISHES   OF  THE  GULF   OF   MAINE 


63 


shore  fishing  grounds,  for  its  European  relative  is  a 
well-known  enemy  of  the  cod,  and  there  is  no  rea- 
son to  suppose  that  the  barn-door  skate  is  less 
voracious.  It  bites  readily  on  almost  any  bait, 
and  is  often  caught  on  hand  and  long  lines  as  well 
as  in  otter  trawls,  and  in  weirs  along  shore. 

Little  is  known  of  the  breeding  habits.  The 
yellowish  or  greenish  brown  egg  cases  are  about 
4%  to  5%  inches  (124-132  mm.)  long  by  2%  to  2% 
(68-72  mm.)  inches  broad,  not  counting  the  horns, 
and  thus  much  larger  than  those  of  any  other 
Gulf  of  Maine  skate.  Females  containing  fully 
formed  egg  capsules  have  been  taken  in  December 
and  January  in  Nova  Scotia  waters,  evidence  that 
the  eggs  are  laid  in  winter.  However,  it  seems 
that  the  young  are  not  hatched  until  late  spring  or 
early  summer,  for  we  have  seen  one,  taken  on 
Nantucket  Shoals  in  July,  so  small  (about  7%  in. 
long)  that  it  could  not  have  been  set  free  long  be- 
fore its  capture. 

General  range. — Atlantic  Coast  of  North  America 
from  the  Banks  of  Newfoundland,  Gulf  of  St. 
Lawrence  and  outer  coast  of  Nova  Scotia  and  the 
Nova  Scotia  Banks  to  North  Carolina.64  It  is 
replaced  in  European  seas  by  a  very  close  ally,  the 
common  skate,  Raja  bails. 

Occurrence  in  the  Gulj  oj  Maine. — This  is  a  com- 
mon fish  in  all  parts  of  our  Gulf,  and  any  very  large 
skate  taken  or  reported  there  is  almost  certain  to 
be  a  "barn-door."  Following  the  coast  around 
from  east  to  west  we  find  it  reported  as  plentiful 
off  the  outer  Nova  Scotia  shore;  it  is  known  from 
St.  Mary  Bay;  is  found  very  generally  though  not 
abundantly  in  the  Bay  of  Fundy  and  in  Passama- 
quoddy  Bay;  is  reported  from  Eastport,  Casco 
Bay,  and  generally  along  the  coast  of  Maine;  is 
known  from  various  localities  in  Massachusetts 
Bay,  where  we  have  seen  many  caught;  and  its 
abundance  on  Georges  Bank  and  on  Nantucket 
shoals  is  illustrated  by  an  average  catch  of  about 
21  per  haul  (about  14  percent  of  all  the  skates 
caught) ,  in  42  trawl  hauls  by  the  Eugene  H,  fishing 
from  Nantucket  Lightship  out  into  the  south 
central  part  of  Georges  Bank  in  late  June  1951. 
In  short,  it  is  to  be  expected  anywhere  within  the 
limits  of  the  Gulf.  Like  most  other  skates,  it  is 
often  taken  in  shoal  water  in  our  Gulf  in  summer; 
seldom  or  never  in  winter.  Huntsman  tells  us 
that  it  comes  to  Passamaquoddy  Bay  from  May 
to  November.    We  once  caught  one  nearly  5  feet 

•*  Doubtfully  reported  from  Florida. 


long  at  Cohasset  in  Massachusetts  Bay  in  less  than 
a  fathom  of  water  in  midsummer;  indeed,  it  is 
often  stranded  on  the  beach.  This  inshore  migra- 
tion, however,  does  not  involve  the  entire  stock, 
witness  its  presence  in  20  to  60  fathoms  on  Georges 
Bank  and  off  Cape  Cod  throughout  the  year,  and 
the  fact  that  it  is  reported  by  fishermen  and  has 
been  trawled  by  vessels  of  the  former  Bureau  of 
Fisheries,  also  by  the  Atlantis,  as  deep  as  100 
fathoms  in  summer.  In  the  warmer  waters  off 
the  southern  coast  of  New  England  it  comes  in- 
shore in  spring  and  autumn,  descending  to  some- 
what deeper  water  for  the  summer. 

Commercial  value. — The  barn-door  skate  is  of 
no  commercial  value  except  as  entering  into  the 
small  landings  of  skates  mentioned  on  page  61. 

Big  skate  Raja  ocellata  Mitchill  1815 

Spotted   skate;   Winter   skate;   Eyed   skate 

Bigelow  and  Schroeder,  1953,  p.  240. 

Garman,  1913,  p.  339,  pi.  29,  fig.  2,  as  Rata  diaphanes. 

Description. — This  skate  looks  very  much  like 
the  little  skate,  but  it  is  larger  and  has  more 
numerous  teeth.  The  front  angle  of  the  disc  is 
much  blunter  than  a  right  angle,  bulging  opposite 
the  eyes,  and  the  tip  of  the  snout  is  rounded. 
The  teeth  are  in  from  72  to  110  series  in  each 
jaw  instead  of  66  series,  or  fewer  as  in  erinacea, 
and  they  are  sharper  in  males  than  in  females. 
The  backs  of  both  sexes  are  rough  with  sharp 
spines  on  the  head,  around  the  eyes,  along  the 
anterior  margins  of  the  pectorals,  over  the  shoul- 
ders, and  on  the  sides  of  the  tail.  The  midline 
of  the  back  behind  the  shoulders  is  almost  always 
free  of  spines  in  adults.  But  we  have  one  speci- 
men, a  female  18  inches  long  taken  near  Jeffreys 
Ledge,  November  1,  1927,  which  bears  a  row  of 
large  spines  along  the  midline  of  back  and  tail 
from  the  shoulder  girdle  to  the  first  dorsal  fin. 
Males,  like  those  of  other  skates,  have  rows  of 
retractile  hooks  on  the  outer  parts  of  the  pectorals. 
The  two  dorsal  fins  are  close  together;  the  outer 
corners  of  the  pectorals  are  bluntly  angular;  the 
claspers  of  adult  males  reach  about  halfway  back 
along  the  tail,  which  occupies  about  half  the 
total  length  of  the  fish. 

Color. — Light  brown  above  with  round  dark 
brown  spots.  As  a  rule  there  is  a  large  white 
eye  spot  with  black  center  near  the  rear  corner 
of  the  pectoral  fin,  and  often  two  smaller  ones 


64 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


4. 


/.'•'; 


vtx 


■ 


:&\% 


& 


l 


gs 


7 


Figure  26. — Big  skate  (Raja  ocellala),  male,  about  36  inches  long.     From  Jordan  and  Evermann.     Drawing  by  H.  L. 

Todd. 


close  to  it.  And  we  have  seen  two  large  speci- 
mens from  Georges  Bank  with  several  of  these 
eye  spots  on  each  side  of  the  disk.  There  is  a 
translucent  or  white  area  on  each  side  of  the 
snout  in  front  of  the  eyes  and  the  lower  surface 
is  white. 

The  eye  spots,  if  present,  serve  to  identify 
this  skate  at  a  glance;  sometimes,  however,  they 
are  lacking,  in  which  case  half-grown  specimens 
so  closely  resemble  the  little  skate  that  recourse 
must  be  had  to  the  number  of  teeth  to  tell  the 
one  from  the  other. 

Size. — This  skate  does  not  mature  until  at 
least  25  to  26  inches  long,  and  grows  to  about 
3%  feet  in  length,  commoidy  from  30  to  34  inches. 
Specimens  32  inches  in  length  are  about  20  inches 
wide. 

Habits. — Big  skates  feed  on  the  same  diet  as 
little  skates  do  (p.  69).  Rock  crabs  and  squid 
are  favorite  prey,  but  they  also  take  annelid 
worms,  amphipods,  shrimps,  and  razor  clams,  and 
they  eat  whatever  small  fish  are  readily  available, 
the  list  at  Woods  Hole  including  smaller  skates, 
eels,  herring,  alewives,  bluebacks,  menhaden, 
smelt,  launce,  cbub  mackerel,  butterfish,  cunners, 
sculpins,  silver  hake,  tomcod,  and  hake.96 

u  From  Vina]  Edwards'  and  Linton's  notes. 


It  is  caught  right  up  to  the  wharves  in  the 
Gulf  of  St.  Lawrence;  often  comes  into  very  shoal 
water  on  sandy  beaches,  and  we  once  caught  an 
adult  male  in  September  in  only  2  or  3  feet  of 
water  in  Nauset  Marsh  on  the  outer  coast  of 
Cape  Cod,  but  few  are  found  shoaler  in  our  Gulf 
than  2  to  4  fathoms.  They  are  much  more  plenti- 
ful at  25  to  35  fathoms  than  deeper,  on  the 
offshore  grounds,  as  appears  from  average  catches, 
of  48  per  haul  at  26  to  35  fathoms,  but  only  11 
per  haul  at  36  to  49  fathoms,  and  none  at  50  to  75 
fathoms,  in  42  trawl  hauls  by  the  Eugene  H,  fishing 
from  Nantucket  Lightship  to  the  south-central 
part  of  Georges  Bank  in  late  June  1951,  and  very 
few  are  caught  deeper  than  about  50  fathoms 
anywhere. 

In  our  Gulf  they  inhabit  about  the  same  range 
of  temperature  as  the  little  skate  does,  i.  e., 
from  68°  or  so,  for  those  along  the  Massachusetts 
coast  in  summer,  down  to  34-36°  in  the  coastal 
belt  as  a  whole  in  winter,  and  to  near  32°  in  the 
Bay  of  Fundy  region,  at  least  in  some  years. 
In  the  southern  side  of  the  Gulf  of  St.  Lawrence 
they  are  found  in  the  icy  bottom  water  on  the 
banks  as  well  as  shoaler,  where  temperatures  rise 
to  61°  (16°  C.)  or  more  in  summer.  Those  living 
the  shoalest  in  the  southern  part  of  their  range 


FISHES   OF  THE  GULF  OF   MAINE 


65 


must  be  exposed  to  temperatures  as  high,  perhaps, 
as  68°  to  70°  at  the  warmest  time  of  the  year. 

Off  the  Atlantic  Coast  of  Nova  Scotia  this  skate 
deposits  its  eggs  from  summer  into  autumn,  and 
probably  through  the  same  season  in  the  Gulf  of 
Maine  for  Scattergood  66  reports  females  with  egg 
capsules  in  Maine  waters  in  September.  And  it 
continues  to  do  so  into  December  and  January  off 
southern  New  England.  Its  egg  cases  are  larger 
than  those  of  the  little  skate,  2^  to  2%  inches  by 
about  1%  inches,  not  counting  the  horns.  The 
length  of  the  period  of  incubation  is  not  known. 

General  range. — Atlantic  Coast  of  North  Amer- 
ica from  northern  North  Carolina  to  the  southern 
side  of  the  Gulf  of  St.  Lawrence,  and  to  the 
southern  part  of  the  Newfoundland  Banks. 

Occurrence  in  the  Gulf  of  Maine. — This,  the 
second  in  size  of  our  skates,  occurs  commonly  all 
around  the  Gulf  of  Maine  from  Nova  Scotia  to 
Cape  Cod.  There  are  many  locality  records  for 
it  for  the  Bay  of  Fundy  as  well  as  from  the  coasts 
of  Maine  and  Massachusetts,  but  so  closely  does 
a  half  or  two-thirds  grown  big  skate  resemble  an 
adult  little  skate  (p.  68)  that  it  is  often  impossible 
to  tell  to  which  species  published  reports  refer. 
It  also  makes  up  so  considerable  a  proportion  of 
the  skate  population  on  Georges  Bank  that  about 
14  percent  of  the  catch  of  skates  made  on  Georges 
Bank  by  one  otter  trawler  in  September  1929,  and 
about  18  percent  (1,116)  of  the  skates  taken  in  42 
trawl  hauls  by  the  Eugene  H,  late  June  1951,  fish- 
ing from  Nantucket  Lightship  to  the  southwestern 
part  of  Georges  Bank,  were  this  species.  But  it 
has  never  been  reported  from  the  deeper  troughs 
of  the  Gulf,  nor  have  we  taken  it  there. 

The  name  "winter  skate"  seems  appropriate 
enough  for  it  along  the  southern  coast  of  New 
England,  for  it  is  only  during  the  cold  season  that 
big  skates  come  close  inshore  near  Woods  Hole. 
And  they  are  said  to  be  taken  in  larger  numbers  in 
winter  than  in  summer  in  the  Massachusetts  Bay 
region  (we  cannot  verify  this).  However,  this  is 
a  misnomer  in  the  cooler  waters  of  the  northern 
part  of  the  Gulf  of  Maine,  for  it  is  common  inshore 
in  Passamaquoddy  Bay  from  May  to  November, 
and  this  probably  applies  to  the  whole  coastline 
east  of  Cape  Elizabeth  to  judge  from  temperature. 

Big  skates  are  taken  on  hook  and  line,  in  weirs, 
and  in  otter  trawls,  but  they  are  of  no  commercial 
value,  except  as  they  form  a  part  of  the  general 

•  Copela  1951,  No.  2,  p.  169. 
210941 — 53 6 


landings  of  skates.     And  they  are  only  a  nuisance 
to  anglers. 

Brier  skate  Raja  eglanteria  Bosc  1802 

Bigelow  and  Schroeder,  1953,  p.  165. 
Garman,  1913,  pi.  23. 

Description. — In  the  brier  skate,  as  in  the  thorny 
skate,  the  midline  of  the  back  and  tail  is  armed 
with  a  continuous  row  of  stout  thorns  from  the 
shoulders  to  the  first  dorsal  fin  near  the  tip  of  the 
tail,  usually  with  1  or  2  in  the  gap  between  the  2 
dorsal  fins.  But  the  thorns  of  this  row  are  not 
much  larger  than  those  along  the  sides  of  the  tail 
(they  are  in  the  thorny  skate),  and  there  are  at 
least  16  thorns  in  the  midrow  along  the  tail  (not 
more  than  9  to  10  in  the  thorny  skate).  There  also 
are  groups  of  large  thorns  opposite  and  behind  the 
eyes,  with  1  to  5  on  each  shoulder  and  1  to  4  rows 
along  either  side  of  the  tail.  Elsewhere  the  upper 
surface  of  the  disc  bears  only  small  sharp  prickles 
(hence  its  name),  most  numerous  on  the  forward 
parts  of  the  pectorals,  over  head  and  snout,  and 
along  the  middle  of  the  back  and  tail  among  the 
larger  thorns.  Thus  it  is  a  much  smoother  species 
than  the  thorny  skate,  and  its  snout  is  more  acute, 
its  outline  being  about  a  right  angle  with  the  mar- 
gins bulging  less  opposite  the  eyes  than  in  any  of 
the  blunter-nosed  skates.  The  outer  corners  of 
the  pectorals  are  distinctly  angular,  and  the 
dorsal  fins  are  separated  by  a  short  gap. 

Color.— Brownish  to  grayish  above;  the  pec- 
torals variously  marked  with  darker  spots  and 
blotches  and  with  more  elongate  bars;  this  last  is 
a  characteristic  feature;  there  is  a  translucent 
space  on  each  side  of  the  snout;  it  is  white  below. 
It  is  most  readily  recognized  by  its  color  pattern, 
with  short  dark  bars  as  well  as  spots,  which  is 
not  shared  by  any  other  Gulf  of  Maine  skate. 

Size. — The  brier  skate  ordinarily  grows  to  a 
length  of  about  2%  feet.  The  largest  on  record 
was  about  37  inches  long. 

General  range.— OS  the  eastern  coast  of  the 
United  States  from  Massachusetts  Bay  to  both 
coasts  of  Florida. 

Occurrence  in  the  Gulf  of  Maine. — This  is  a 
southern  species,  uncommon  even  as  far  north  as 
Woods  Hole  and  decidedly  rare  in  the  Gulf  of 
Maine.  It  has  been  recorded  once  from  Glouces- 
ter, its  most  northerly  outpost,  and  also  from 


66 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


Figure  27. — Brier  skate  (Raja  eglanteria),  female,  about  29  inches  long,  Woods  Hole,  Massachusetts. 

Schroeder.     Drawing  by  E.  N.  Fischer. 


From  Bigelow  and 


Provincetown.  Two  specimens  were  taken  on 
Nantucket  Shoals  near  Round  Shoal  buoy  by  the 
Halcyon,  one  in  July,  the  other  in  September,  in 
1924. 

Leopard  skate  Raja  garmani  Whitley  1939 

FiOSETTED    SKATE 

Bigelow  and  Schroeder,  1953,  p.  200. 
Garman,  1913,  pi.  18,  fig.  2. 

Description. — The  conspicuous  dark  rosettes  on 
the  upper  surface  make  this  skate  recognizable  at 
a  glance,  since  no  other  skate  of  the  western 
Atlantic  is  marked  in  this  way.  And  its  tail  is 
longer  relatively  than  that  of  any  other  Gulf  of 
Maine  skate. 

The  disc  is  considerably  blunter  in  front  than 
a  right  angle,  with  anterior  margins  bulging 
rather  conspicuously  a  little  anterior  to  the  level 
of  the  eyes;  the  outer  corners  of  the  pectorals  are 
very  broadly  rounded ;  the  tad  measured  from  the 


center  of  the  cloaca  to  the  tip  is  about  1.5  times  as 
long  as  the  body  from  tip  of  snout  to  center  of 
cloaca;  and  there  is  a  definite  gap  with  one  or 
two  thorns  between  the  two  dorsal  fins.67 

There  are  thorns  along  the  ridge  of  the  snout; 
a  row  around  the  inner  and  posterior  margins  of 
the  eyes  with  a  few  in  the  space  between  the  latter; 
a  group  on  each  shoulder;  and  one  row  along  the 
midbelt  of  the  back  and  tad  in  young  specimens, 
increasing  to  2  to  6  irregular  rows  ia  large  ones. 
In  young  specimens  the  skin  of  the  disc,  as  a  whole, 
and  of  the  tail,  is  also  rough  with  small  prickles,  but, 
most  of  these  are  lost  with  growth,  leaving  large 
specimens  mostly  naked  except  for  the  thorns. 
The  lower  surface  is  smooth. 

There  are  46  to  52  series  of  teeth  in  the  upper 
jaw,  a  few  less  in  the  lower,  and  those  of  adult 
males  are  only  a  little  sharper  than  those  of 
females. 

Color. — The  upper  side  is  pale  buff  or  brown, 


87  Garman's  illustration  is  of  an  abnormal  specimen  with  three  dorsal  flna. 


FISHES   OF  THE  GULF  OF   MAINE 


67 


Figure  28. — Leopard  skate  (Raja  garmani),  female,  16  inches  long,  offing  of  Montauk  Point,  New  York.     From  Bigelow 

and  Schroeder.     Drawing  by  E.  N.  Fischer. 


freckled  with  small  spots,  darker  or  lighter,  and 
conspicuously  marked  with  dark  rosettes,  each 
consisting  of  a  group  of  6  or  more  dark  brown  or 
black  spots  surrounding  a  dark  central  spot.  The 
lower  surface  is  white  or  pale  yellow. 

Size. — This  is  one  of  the  smaller  skates,  males 
maturing  when  only  about  16  inches  long. 

General  range. — Outer  part  of  the  continental 
shelf  and  upper  part  of  the  continental  slope  from 
southern  Florida  to  the  offing  of  Nantucket,  in 
depths  of  30  to  300  fathoms. 

Occurrence  in  the  Qulj  of  Maine. — Our  only 
reason  for  mentioning  this  species  is  that  one 
specimen  was  trawled  by  the  Albatross  HI,  May 
14,  1950,  at  52  fathoms  southeast  of  Nantucket 
Lightship  Gat.  40°05'  N.,  long.  69°22'  W.). 
And  this  is  probably  close  to  the  eastern  boundary 
of  its  range,  for  it  has  never  been  reported  among 
the  other  skates  that  are  trawled  in  abundance 
along  the  seaward  slopes  of  Georges  and  of  the 
Nova  Scotia  Banks.  But  it  is  one  of  the  most 
plentiful  of  skates  offshore  to  the  westward,  along 
southern  New  England. 


Little  skate  Raja  erinacea  Mitchill  1825 

Common    skate;    Summer    skate;    Hedgehog 
skate;  Tobacco  box 

Bigelow  and  Schroeder,  1953,  p.  176. 
Garman,  1913,  pi.  20. 

Description. — The  most  distinctive  characters  of 
grown  specimens  are  their  small  size,  absence  of 
thorns  along  the  midline  of  the  back  (except  in 
the  young)  and  blunt  nose. 

The  anterior  angle  of  the  disc  is  blunter  than 
a  right  angle  and  the  tip  of  the  snout  is  rounded, 
with  the  margins  bulging  opposite  the  eyes.  The 
teeth  are  in  only  about  38  to  66  series.  Females 
have  thorns  scattered  generally  over  the  upper 
surface;  these  are  especially  prominent  on  head, 
snout,  shoulders,  and  sides  of  tail.  Ordinarily 
there  are  no  spines  on  the  midline,  back  of  the 
shoulder  girdle;  but  we  found  one  fish,  13 K  inches 
long,  among  the  many  we  have  observed,  with 
a  median  row  of  spines  extending  from  the 
shoulder  girdle  to  the  first  dorsal  fin  near  the 


68 


FISHERY   BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


Figure  29. — Little  skate  (Raja  erinacea).  A,  male,  20  inches  long,  Boston  Harbor;  B,  female,  17%  inches  long,  Mystic 
Connecticut;  C,  side  view,  end  of  tail  of  same,  about  0.6  times  natural  size.  From  Bigelow  and  Schroeder.  Drawings 
by  E.  N.  Fischer. 


end  of  the  tail,  and  this  is  true  of  newly  hatched 
specimens  in  general.  Males  are  less  spiny,  but 
the  spines  on  tail,  shoulders,  and  along  either 
side  of  the  back  ridge  are  noticeably  strong  in 
both  sexes.  The  two  dorsal  fins  are  close  together; 
the  tail  is  about  half  the  total  length.  Large 
ones  closely  resemble  small  specimens  of  the  big 
skate  (R.  ocellata,  p.  63)  that  may  chance  to 
lack  the  ocellar  spots  with  which  that  species 
usually  is  marked.  A  count  of  the  teeth  is  then 
the  only  sure  clue  to  the  identity  of  the  specimen 
in  hand. 

Color. — Grayish  to  dark  brown  above,  or 
clouded  light  and  dark  brown,  paler  at  the  edges 
of  the  pectoral  fins;  usually  with  many  small  round 
darker  spots;  white  or  grayish  below. 

Size. — Ordinarily  up  to  16  to  20  inches  long; 


the  maximum  recorded  length  is  21  inches  (53 
cm.);  they  weigh  about  %  to  1  pound  at  16  to  17 
inches  and  anywhere  from  1%  to  2  pounds  at  18 
inches.  Females  mature  sexually  when  12%-17 
inches  (32-43  cm.)  long,  males  at  about  14  to 
17K  inches.68 

Habits. — It  is  common  knowledge  that  this 
skate,  like  others,  is  most  abundant  on  sandy 
or  pebbly  bottom;  but  they  are  likewise  found 
on  mud  and  we  have  seen  them  lying  on  ledges 
at  times. 

The  usual  depth  range  is  from  close  to  tide  line 
down  to  75  fathoms  or  so.  Many  even  follow 
up  the  shelving  bottoms  of  our  beaches  until  they 


88  Information  supplied  by  Dr.  Daniel  Merrimau,  Dr.  Y.  H.  Olsen,  the 
Misses  S.  B.  Wheatland  and  L.  H.  Calhoun,  who  have  made  a  detailed 
study  of  the  littlo  skate  in  southern  New  England  waters. 


FISHES  OF  THE  GULF  OF   MAINE 


69 


strand.  And  tho  bulk  of  the  population  hold 
to  depths  of  less  than  40  to  50  fathoms,  as 
appears  from  average  catches,  per  haul,  of  100 
at  26  to  35  fathoms,  and  95  at  36  to  49  fathoms, 
but  only  12  at  50  to  75  fathoms,  in  42  hauls  by 
the  Eugene  H,  fishing  from  Nantucket  Lightship 
to  the  southcentral  part  of  Georges  Bank,  in  late 
June  1951.  Fifty  fathoms  (off  the  Bay  of  Fundy) 
is,  in  fact,  the  deepest  that  positively  identified 
specimens  are  known,  in  the  inner  parts  of  our 
Gulf;  80  fathoms  off  southern  New  England.69 

The  little  skate  tolerates  a  wide  range  of 
temperature,  being  found  in  water  as  warm  as 
68-70°  in  summer,  while  they  are  exposed  to 
temperatures  close  to  32°  in  the  Bay  of  Fundy 
in  some  winters,  unless  they  move  out,  and 
deeper  there  than  seems  likely.  In  the  southern 
side  of  the  Gulf  of  St.  Lawrence,  writes  Hunts- 
man,70 they  are  found  in  the  intermediate  zone 
between  the  icy  cold  waters  of  the  banks  and  the 
surface  stratum,  which  last  warms  to  61°  (16° 
C.)  or  higher  in  summer. 

They  have  never  been  reported,  to  our  knowl- 
edge, where  the  water  is  appreciably  brackish. 

Hermit  and  other  crabs,  shrimps,  worms, 
amphipods,  ascidians  ("sea  squirts"),  bivalve 
mollusks,  squid,  small  fishes,  and  even  such  tiny 
objects  as  copepods  have  been  found  in  their 
stomachs.  Probably  crabs  loom  largest  in  their 
diet,  for  more  than  29  percent  of  the  skates 
opened  by  Field  at  Woods  Hole,  contained  them; 
15  percent  had  bottom-dwelling  shrimps  (Crago); 
and  6  percent  had  eaten  squid.  In  Long  Island 
Sound,  however,  amphipods  (Leptocheirus)  are 
the  dominant  item  in  their  diet,  forming  from 
one- third  to  one-half  of  the  stomach  contents  at 
all  seasons  of  the  year.71  Launce,  alewives,  her- 
ring, cunners,  silversides,  tomcod,  silver  hake, 
have  all  been  found  in  their  stomachs,  and  they 
bite  a  baited  hook  readily,  affording  amusement 
to  vacationists. 

The  spawning  habits  of  the  little  skate  have  not 
been  followed  in  the  Gulf  of  Maine.  Studies,  at 
the  Bingham  Oceanographic  Laboratory,  however, 
suggest  that  they  ordinarily  deposit  their  eggs  in 
water  not  deeper  than  15  fathoms  and  on  a  sandy 
bottom.  It  appears  from  anatomical  examination 
of  the  sexual  organs  of  the  mature  females  that 

n  Seventeen  that  we  saw  trawled  on  the  Albatross  III,  May  1950. 
"  Trans.  Roy.  Soc.  Canada,  Ser.  3,  vol.  12,  sec.,  4, 1918,  p.  63. 
71  Information  from  Dr.  Daniel  Merriman,  Dr.  Y.  H.  Olsen,  and  the 
Misses  S.  B.  Wheatland  and  L.  H.  Calhoun. 


copulation  may  take  place  at  any  time  throughout 
the  year,  and  frequently.  Observations,  too,  on 
skates  kept  in  aquaria  have  shown  that  the  eggs 
are  laid  in  pairs  at  intervals  of  from  five  days  to 
several  weeks;  also  that  they  are  usually  buried  in 
sand,  at  least  partially.72  The  eggs  have  been 
taken  off  Southern  New  England,  in  fish  traps 
and  dredges  in  a  few  fathoms  of  water  in  abun- 
dance from  July  through  September. 

Examination  of  large  numbers  of  females  has 
shown  that  eggs  are  laid  there  throughout  the 
year.  And  there  is  no  reason  to  doubt  that  this 
is  the  case  to  the  north  and  east  of  Cape  Cod  as 
well.  Aquarium  experiments  have  also  shown 
that  eggs  laid  in  the  period,  May-July,  hatched 
between  the  end  of  November  and  the  beginning 
of  January,  i.  e.,  after  5  to  6  months.  But  the 
incubation  period  is  likely  to  be  somewhat  longer 
for  spring-summer  laid  eggs  in  nature  because  of 
somewhat  lower  temperatures;  and  considerably 
longer  for  eggs  laid  in  autumn  and  early  winter. 

The  eggs  measure  about  \){  to  1%  inches  by 
about  2%  to  2%  inches,  not  counting  the  horns, 
and  the  great  majority  of  the  empty  skate  eggs 
that  are  washed  up  on  the  beaches  of  our  Gulf 
belong  to  this  species.  The  young  skate,  which 
emerges  through  a  transverse  opening  at  the  edge 
of  the  egg  case  at  the  end  that  has  the  longer  pair 
of  horns,  is  about  3%  to  4  inches  long  at  hatching; 
its  abdomen  is  still  swollen  with  yolk,  and  its 
tail  terminates  in  a  whiplash-like  extension  that 
disappears  within  a  few  days.  Huntsman's  ob- 
servations suggest  that  young  hatched  near  the 
head  of  the  Bay  of  Fundy  descend  to  deeper 
water  the  first  winter,  and  this  probably  applies 
to  the  Gulf  of  Maine  as  a  whole. 

It  appears  from  information  of  various  sorts 
that  a  little  skate  8  inches  (20  cm.)  long  may  be  1 
to  1%  years  old;  one  of  11%  to  12  inches  (30  cm.) 
2  to  3  years;  one  of  15%  to  16  inches  (40  cm.)  3 
to  4  years;  one  of  19%  to  20  inches  6  to  8  years  old. 
And  the  mortality  rate  appears  to  be  very  high 
after  five  years,  for  very  few  of  those  taken  are 
longer  than  about  18  to  19  inches.73 

General  range. — Atlantic  coast  of  America; 
southern  side  of  the  Gulf  of  St.  Lawrence  and 


,3  This  summary  of  the  breeding  habits  is  based  on  extensive  information 
supplied  by  Dr.  Daniel  Merriman,  Dr.  Y.  H.  Olsen,  and  the  Misses  S.  B. 
Wheatland  and  L.  H.  Calhound. 

'3  Information  from  Dr.  Daniel  Merriamn,  Dr.  Y.  H.  Olsen,  and  the 
Misses  S.  B.  Wheatland  and  L.  H.  Calhoun. 


70 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


northern  Nova  Scotia  to  Virginia,  in  coastal  waters 
and  on  the  shoaler  of  the  offshore  banks. 

Occurrence  in  the  Gulf  of  Maine. — This,  the 
smallest  of  our  skates,  is  the  commonest  and  the 
most  familiar  from  its  habit  of  coming  up  into 
very  shoal  water  in  summer  and  of  stranding  on 
the  beaches,  where  dried  skate  carcasses  are  often 
to  be  seen.  It  occurs  all  along  the  coast  in  the 
southern  side  of  the  Gulf  of  St.  Lawrence  and 
along  outer  Nova  Scotia;  is  very  abundant  both  on 
the  New  Brunswick  and  on  the  Nova  Scotia  sides 
of  the  Bay  of  Fundy,  and  is  to  be  caught  every- 
where and  anywhere  along  the  coasts  of  Maine 
and  of  Massachusetts ;  far  more  commonly,  indeed, 
than  one  might  suspect  from  the  few  definite 
records  that  have  found  their  way  into  scientific 
literature. 

An  average  catch  of  about  88  per  haul  (about 
60  percent  of  all  the  skates  taken)  in  42  trawl 
hauls  by  the  Eugene  H,  in  late  June  1951,  fishing 
eastward  from  Nantucket  Lightship  suggests  that 
this  is  the  most  plentiful  skate  on  the  south- 
western part  of  Georges  Bank  and  on  the  Nan- 
tucket grounds.  But  it  seems  to  be  far  less 
numerous  on  the  northeastern  part  of  the  Bank, 
if  it  is  present  there  at  all;  at  least  we  failed  to 
find  a  single  one,  among  495  skates  of  other  kinds 
caught  there  in  37  hauls  by  the  otter  trawler 
Kingfisher  in  September  1929.  And  we  have 
never  found  it  (nor  has  it  been  reported)  in  the 
deeper  basins  and  troughs  of  our  Gulf,  probably 
because  it  is  restricted  in  general  to  depths  less 
than  40  to  50  fathoms  (p.  69). 

In  our  Gulf  many  of  the  little  skates  appear  to 
carry  out  an  irregular  migration  into  shoal  water 
in  April  and  May,  where  they  remain  throughout 
the  summer,  autumn,  and  early  winter,  to  retire 
again  to  somewhat  deeper,  hence,  warmer  water 
in  December  or  January.  Its  migration  schedule 
appears  to  be  more  complex  in  Long  Island  Sound 
waters  where  summer  temperatures  are  higher; 
i.  e.,  inshore  in  spring,  offshore  in  mid-  or  late 
summer,  inshore  again  in  late  autumn  and  offshore 
again  in  midwinter.74  Doubtless  little  skates 
breed  throughout  the  shoaler  parts  of  the  Gulf, 
and  on  the  offshore  banks. 

They  are  of  no  commercial  importance  in  our 
Gulf  except  as  they  form  a  part  of  the  landings  of 
trash  fish. 


"  Information  from  Dr.  Daniel  Merriman,  Dr.  Y.  H.  Olsen,  the  Misses 
S.  B.  Wheatland  and  L.  H.  Calhoun  of  the  Bingham  Oceanographic  Lab- 
oratory. 


Smooth-tailed  or  prickly  skate  Raja  senta 
Garman  1885 

Bigelow  and  Schroeder,  1953,  p.  264. 
Garman,  1913,  pi.  25,  fig.  1. 

Description. — By  the  time  this  skate  has  grown 
to  one-fourth  its  adult  size  it  is  made  recognizable 
by  the  fact  that  the  middorsal  line  of  thorns  runs 
back  only  to  about  the  middle  of  the  tail,  where  the 
thorns  so  dwindle  in  size  that  they  are  not  dis- 
tinguishable from  the  tiny  prickles  with  which  the 
tail  is  clothed,  generally.  Newly  hatched  speci- 
mens in  which  this  character  is  not  yet  established 
are  separable  from  all  other  Gulf  of  Maine  skates 
by  the  color  pattern  of  the  tail,  which  has  two  pale 
crossbars,  each  outlined  in  front  and  behind  by  a 
dark  band  or  blotch. 

There  is  a  single  row  of  16  or  more  medium- 
sized  to  large  thorns  along  the  midline  of  the  back, 
spaced  irregularly,  and  usually  about  20  to  30 
along  the  anterior  one-half  or  so  of  the  tail ;  about 
10  to  13  around  the  inner  ridge  of  each  eye;  and 
3  to  5  on  each  shoulder.  Immature  specimens  of 
both  sexes  are  also  closely  and  uniformly  rough- 
ened with  small  prickles  over  the  disc  as  a  whole, 
on  the  pelvics  and  on  the  upper  side  of  the  tail. 
But  irregular  bare  areas  develop  on  the  shoulders 
and  around  the  outer  parts  of  the  pelvics  of  females 
as  they  approach  maturity  while  mature  males 
lose  the  prickles  from  the  central  part  of  the  disc 
as  a  whole,  but  develop  a  few  thorns  on  the  mid- 
ridge  of  the  snout  besides  larger  thorns  over  a 
roughly  triangular  area  on  either  side  of  the  head 
abreast  of  the  eyes  and  farther  forward.  They 
also  develop  two  rows  of  the  usual  curved  sexual 
spines  on  either  side  on  the  pectorals,  about  13 
to  14  per  row.  The  lower  surface  of  the  disc  is 
smooth,  except  that  a  few  prickles  develop,  with 
growth,  along  the  margins  near  the  snout 

The  lower  surface  of  the  tail  as  a  whole  is 
prickly  on  females  and  on  immature  males,  but 
tends  to  become  smooth  on  males  by  the  time 
they  mature  sexually. 

The  anterior  angle  of  the  snout  is  a  little  more 
obtuse  than  a  right  angle  (about  110°);  the  tip  of 
the  snout  is  sharper  than  in  either  the  big  skate, 
the  little  skate,  or  the  thorny  skate.  There  are  38 
to  40  series  of  teeth  in  the  upper  jaw,  36  to  38 
series  in  the  lower  jaw;  those  of  females  are  low, 
with  only  faintly  indicated  points,  but  those  of 
mature  males  are  longer,  sharper,  recurved,  and 


FISHES    OF  THE   GULF   OF    MAINE 


71 


Figure  30. — Smooth-tailed  or  prickly  skate  {Raja  senta),  male,  about  20%  inches  long,  Emerald  Bank,  Nova  Scotia. 

From  Bigelow  and  Schroeder.     Drawing  by  E.  N.  Fischer. 


spaced  more  loosely.  There  is  no  free  space 
between  the  two  dorsal  fins.  The  disc  is  a  little 
broader  than  long  (1.2  to  1.3  times);  the  tail 
occupies  about  one-half  of  the  total  length. 

Color. — -The  upper  side,  including  the  tail,  is  pale 
brown,  with  many  obscure  darker  spots.  Newly 
hatched  specimens  are  also  marked  on  the  tail  with 
two  pale  cross  bars,  each  outlined  by  a  darker  cross 
bar  or  blotch  in  front  of  it  and  one  behind,  but 
these  bars  disappear  with  growth.  The  lower 
surface  is  white,  either  plain  or  with  a  few  dusky 
blotches.  Sometimes  the  rear  part  of  the  tail  is 
uniformly  dark  below. 

Size.- — -A  male  about  20  inches  (515  mm.)  long 
that  we  have  seen  seems  to  be  sexually  mature. 
The  largest  specimen  of  which  we  have  record 
was  24  inches  long. 

Habits. — -This  skate  appears  equally  at  home 
on  the  soft  mud  and  clay  bottoms  of  the  deeper 
basins  of  the  Gulf  and  on  the  sand,  broken  shells, 
gravel  and  pebbles  of  the  offshore  fishing  banks. 
Nothing  is  known  of  its  diet.  Egg  cases,  appar- 
ently of  this  species,  have  been  trawled  in  deep 
water  (82-164  fathoms),  in  the  estuary  of  the  St. 


Lawrence  River  in  July  and  August;  probably 
they  are  laid  in  summer  in  the  Gulf  of  Maine,  as 
well. 

General  range. — Atlantic  shelf  of  North  America 
from  the  offing  of  Charleston,  S.  C,  to  the  Nova 
Scotia  Banks  and  Gulf  of  St.  Lawrence,  a  few 
reaching  the  southern  part  of  the  Newfoundland 
Banks;  mostly  in  depths  greater  than  about  40 
to  50  fathoms.  The  deepest  record  for  it  is  478 
fathoms  off  South  Carolina. 

Occurrence  in  the  Gulf  of  Maine. — This  skate, 
once  considered  a  rare  species,  is  now  known  to 
occur  generally  throughout  the  western  side  of  the 
Gulf  wherever  the  water  is  more  than  45  to  50 
fathoms  deep,  25  fathoms  being  our  shoalest 
record  for  it  and  on  the  offshore  Banks  as  well. 
We  have  trawled  it  on  several  occasions  in  the 
deep  trough  west  of  Jeffreys  Ledge ;  in  deep  water 
(80-100  fathoms)  near  Cashes  Ledge;  also  in  the 
basin  east  and  southeast  of  Cape  Cod.  And, 
being  known  from  the  southeastern  slope  of 
Browns  Bank,  it  is  to  be  expected  generally  in  the 
eastern  side  of  the  Gulf,  as  well  as  in  the  western, 
at  the  proper  depth.     It  is  widespread  on  Georges 


72 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


Bank  also,  but  is  far  less  plentiful  there  than  other 
skates,  to  judge  from  the  fact  that  trawl  hauls  in 
September  1929  brought  in  only  37  of  them,  and 
that  we  counted  only  8,  from  42  hauls  on  the 
Eugene  H;  in  June  1951,  fishing  from  Nantucket 
Lightship  to  the  south  central  part  of  Georges. 
We  have  trawled  it  at  50  to  250  fathoms  off 
southern  New  England.  To  the  eastward  and 
northward,  it  is  recorded  on  La  Have  and  Emerald 
Banks  at  50  to  100  fathoms,  and  in  the  Gulf  of  St. 
Lawrence  at  82  to  178  fathoms. 

Thorny  skate  Raja  radiata  Donovan  1807 

Starry  skate76 

Bigelow  and  Schroeder,  1953,  p.  255." 
Garman,  1913,  pi.  21,  fig.  2.™ 

Description. — The  thorny  skate  can  be  identified 
at  a  glance  among  skates  of  the  Gulf  of  Maine  by 
the  fact  that  the  row  of  thorns  with  which  the 
midline  of  back  and  tail  is  armed  are  not  only 
large  and  conspicuous,  but  do  not  number  more 
than  19  at  most  from  the  nape  back  along  the  tail. 
There  are  also  2  or  3  large  thorns  on  each  shoulder; 
and  one  in  front  of  each  eye  and  one  behind  it ;  one 
close  to  the  inner  end  of  each  spiracle;  and  other 
smaller  thorns  scattered  on  snout,  pectoral  fins, 
and  tail.  The  bases  of  the  thorns  on  the  pectorals 
are  star-shaped,  a  very  distinctive  character; 
those  of  the  still  larger  thorns  along  the  midline  of 
the  back  are  oval.  Adult  males  have  2  rows  of 
hooked,  erectile  thorns  near  the  outer  corners  of 
the  pectorals. 

The  anterior  angle  of  the  disc  is  considerably 
more  obtuse  than  a  right  angle  (110-140°),  and  the 
tip  of  the  snout  is  blunt  with  the  margins  bulging 
somewhat  a  little  in  front  of  the  level  of  the  eyes; 
the  outer  corners  of  the  pectorals  are  less  broadly 
rounded  than  in  either  the  little  skate  or  the  big 
skate;  and  the  two  dorsal  fins  may  either  be 
joined  at  the  base  or  be  separated  by  a  short 
space.  There  are  36  to  46  series  of  teeth  in  each 
jaw,  those  of  females  and  of  young  males  with  low 
cusps   that  are  worn  nearly  smooth   along   the 

11  When  the  first  edition  of  this  book  appeared,  it  was  an  open  question 
whether  the  thorny  skate  of  American  waters  (named  R.  scabrata  by  Garman 
1913)  was  identical  with  the  thorny  skate  of  northern  Europe  (R.  radiata 
Donovan,  1807).  Our  subsequent  comparison  of  American  specimens  with 
others  from  Greenland  and  Norway  has  convinced  us  that  they  all  belong  to 
the  one  species,  which  must  be  called  by  the  older  of  the  two  scientific  names. 

"  Figure  1  of  Garman's  plate  21  is  not  of  a  thorny  skate,  as  it  is  named  to  the 
accompanying  caption,  but  is  of  a  small  specimen  of  the  big  skate  that  wc  have 
examined. 


older  rows;  those  of  mature  males  a  little  sharper 
and  spaced  a  little  more  widely. 

Color. — Brown  above,  either  uniform  or  slightly 
clouded,  or  spotted  with  darker,  small  specimens 
more  conspicuously  so  than  larger.  Sometimes 
there  is  a  white  spot  beside  each  eye,  one  on  either 
side  abreast  of  the  nape,  and  another  on  each  side 
on  the  rear  part  of  the  disc.  The  lower  side  is 
white,  sometimes  with  irregular  sooty  or  brownish 
blotches.  Garman  mentions  a  partial  albino, 
white  above  with  a  few  reddish  brown  and  brown 
spots. 

Size. — The  thorny  skate  is  about  4  inches  (100 
mm.)  long  from  snout  to  first  dorsal  fin  at  hatch- 
ing. The  largest  specimens  so  far  recorded  from 
American  waters  have  been  about  40  inches  for 
the  Nova  Scotia  Banks,  35%  inches  for  Georges 
Bank,  and  about  31  inches  for  Massachusetts 
Bay.  But  some  males  may  mature  when  only  21 
to  22  inches  long.  One  32  inches  long  is  about  23 
inches  wide. 

Habits. — The  thorny  skate  is  a  cool  water  fish, 
at  home  in  temperatures  from  about  50°  or  so  down 
nearly  to  the  freezing  point  of  salt  water.  It  is 
also  restricted  in  general  to  depths  greater  than 
about  10  fathoms,  even  in  the  northernmost  part 
of  its  range.  In  the  Gulf  of  St.  Lawrence  it  lives 
indifferently  on  the  ice  cold  banks  and  in  the 
warmer  water  on  the  bottom  of  the  deep  Lauren- 
tian  Channel.  Average  catches  of  1  per  haul  at 
26  to  35  fathoms,  22  per  haul  at  36  to  49  fathoms, 
and  5  per  haul  at  50  to  75  fathoms,  in  42  trawl 
hauls,  by  the  Eugene  H  fishing  from  Nantucket 
Lightship,  the  central  part  of  Georges  Bank, 
June  1951,  suggest  a  rather  definite  preference  for 
the  intermediate  depth  zone,  perhaps  because  of 
the  food  supply.  But  thorny  skates  have  been 
taken  at  many  stations,  also,  down  to  336  fathoms 
off  the  American  coast,  and  as  deep  as  459  fathoms 
near  Spitzbergen. 

The  stomachs  of  thorny  skates  caught  on 
Georges  Bank  contained  shrimps,  spider  crabs, 
anemones,  hydroids,  and  fish  digested  past 
identification. 

The  egg  cases  vary  considerably  in  size,  prob- 
ably depending  on  the  size  of  the  parent  fish. 
One  from  a  fish  32  inches  long,  taken  on  Georges 
Bank,  measured  3  by  2%  inches  exclusive  of  the 
horns.  Others  that  have  been  measured  from  the 
Nova  Scotia  Banks  ranged  from  3  to  3%  inches  in 
length.    They  are  flat  on  one  side,  strongly  convex 


FISHES   OF  THE  GULF   OF   MAINE 


73 


Figure  31. — Thorny  skate  (Raja  radiala),  female,  about  31%  inches  long.     After  Garman. 


on  the  other,  and  are  rough  with  narrow  cross- 
ridges.  A  mass  of  delicate  fibrils,  matted  to- 
gether, extends  along  each  of  the  longer  sides  and 
partly  over  the  surfaces  also.  And  each  horn  ends 
in  a  slender  fibril. 

General  range. — The  thorny  skate  is  known  on 
both  sides  of  the  northern  Atlantic.  In  the  east 
its  range  extends  from  the  White  Sea  and  Barents 
Sea  to  the  North  Sea,  Dutch  coast,  and  western 
part  of  the  Baltic;77  in  the  west  from  West  Green- 
land, Hudson  Bay,  Atlantic  coast  of  Labrador, 
east  and  south  coasts  of  Newfoundland,  Grand 
Banks,  Gulf  of  St.  Lawrence  and  outer  coast  of 
Nova  Scotia  with  the  off-lying  fishing  grounds,  to 
the  Gulf  of  Maine,  and  thence  westward  and  south- 
ward along  the  edge  of  the  continental  shelf  to  the 

77  Doubtfully  reported  from  Belgium  and  the  Bay  of  Biscay. 


offing  of  New  York;  and  as  a  stray  to  the  offing 
of  Charleston,  S.  C.78 

Occurrence  in  the  Gulf  of  Maine. — The  thorny 
skate  is  not  often  seen  close  inshore  along  our  coast, 
being  restricted  in  general  to  moderately  deep  water 
(p.  72).  But  it  is  now  known  to  be  generally  dis- 
tributed in  the  deeper  waters  of  the  Gulf.  Thus 
it  is  frequently  taken  on  the  New  Brunswick  side 
of  the  Bay  of  Fundy  in  depths  of  10  fathoms  or 
deeper,  in  20  to  30  fathoms  in  St.  Mary  Bay  on 
the  Nova  Scotia  side.  It  has  been  recorded  from 
Casco  Bay;  from  Ipswicb  Bay,  off  Gloucester, 
Salem  and  Nahant,  and  off  Provincetown ;  and 
we  have  taken  it  ourselves  in  numerous  places  in 
the  Gulf  at  14  fathoms  and  deeper,  including  the 

"  One  taken  in  lat.  33°10'  N.,  long.  77°25'  W.,  in  74  fathoms,  by  the  Albatross 
III  is  in  the  Museum  of  Comparative  Zoology. 


74 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


vicinity  of  Mount  Desert;  PlattsBank;  and  in  the 
bottoms  of  the  deep  troughs.  It  has  also  been 
trawled  at  many  stations  on  Georges  Bank,  like- 
wise along  the  upper  part  of  the  continental  slope 
off  southern  New  England,  down  to  336  fathoms. 
There  is  nothing  in  the  available  record  to  sug- 
gest that  it  carries  out  any  regular  migrations, 
whether  in  or  offshore,  or  along  the  coast.  And  it  is 
more  catholic  in  respect  to  its  choice  of  bottom 
than  some  other  skates,  for  while  it  is  most  plenti- 
ful on  the  good  fishing  grounds  of  sand,  gravel, 
and  broken  shells,  we  have  taken  it  at  many  sta- 
tions in  the  Gulf  on  soft  mud.  And  it  is  one  of  the 
most  plentiful  of  Gulf  of  Maine  skates  at  appro- 
priate depths.  Thus  325  were  caught  in  37  trawl 
hauls  on  the  northeastern  part  of  Georges  Bank  on 
one  trip  in  1929;  again,  in  June  1951,  we  counted 


432,  from  42  trawl  hauls  (7  percent  of  the  total 
catch  of  skates),  on  the  Eugene  H  fishing  from 
Nantucket  Lightship  to  the  south  central  part  of 
Georges.  We  once  caught  12  in  the  western  side 
of  the  Gulf  in  a  beam  trawl  only  8  feet  across  the 
mouth  in  30  minutes;  and  we  have  taken  1  to  100 
of  them  in  26  hauls  with  larger  trawls,  between 
Mount  Desert  Island  and  Massachusetts  Bay. 

Females  containing  eggs  about  ready  to  be  laid, 
and  deposited  eggs  in  various  stages  of  incubation, 
have  been  taken  in  Nova  Scotian  waters  or  in  the 
Gulf  of  Maine,  in  April,  June,  July,  and  September, 
and  they  are  to  be  expected  there  in  winter  as  well, 
having  been  reported  in  January  and  February 
off  Norway,  and  from  February  to  June  in  Scottish 
waters. 


THE  WHIP-TAILED  STING  RAYS.     FAMILY  DASYATIDAE 


The  whip-tailed  sting  rays,  like  the  skates,  are 
disc-like  in  form,  very  thin  toward  the  outer  edges, 
with  the  anterior  parts  of  the  pectoral  fins  fused 
with  the  sides  of  the  head,  and  with  the  eyes  and 
spiracles  on  the  upper  surface.  Their  pelvic  fins, 
however,  have  convex  outer  edges,  not  concave 
as  are  those  of  the  skates.  They  have  no  dorsal 
fin.  Their  tails  are  long  and  whiplash-like  to- 
ward the  tip  and  armed,  in  most  of  them  with  one 
to  several  sawedged,  venomous  spines  on  the 
upper  side.  Their  teeth  are  small  and  in  many 
series,  closely  crowded  in  bands  along  the 
jaws.  The  upper  surface  of  disc  and  tail  is 
smooth  in  some  of  them,  variously  roughened 
with  tubercles,  thorns  or  prickles  in  others. 
They  do  not  lay  eggs  as  the  skates  do,  but  bear 
"living"  young  (p.  57).  And  the  young  resemble 
their  parents  closely  when  born.  Four  species 
are  known  along  the  Middle  and  South  Atlantic 
States,  but  only  one  of  them  reaches  the  Gulf  of 
Maine,  and  then  only  as  a  stray.  Should  any 
long-tailed  sting  ray  be  picked  up  within  the 
limits  of  the  Gulf  that  does  not  fit  the  following 
description,  its  captor  is  referred  to  Bigelow  and 
Schroeder,  1953,79  for  its  identification. 

»  Fishes  Western  North  Atlantic.  Pt.  2.  Mem.  1,  Sears  Foundation,  19S3. 


Sting  ray  Dasyatis  centroura  (Mitchill)  181580 

Stingaree;  Clam  cracker 

Bigelow  and  Schroeder,  1953,  p.  352. 

Garman,  1913,  pi.  33,  figs.  1,  2,  aa  Dasybatus  marinus. 

Description. — The  most  distinctive  features  of 
this  sting  ray,  among  other  Gulf  of  Maine  fishes, 
are  its  very  long,  whiplash-like  tail  without  dorsal 
fins,  and  the  long,  sawedged  spine  or  spines  with 
which  the  upper  side  of  its  tail  is  armed.  The  disc 
is  rhomboid,  about  1%  to  1%  times  as  broad  as  it 
is  long;  the  anterior  angle  is  much  blunter  than  a 
right  angle  (130-140°);  and  the  tip  of  the  snout 
projects  very  little  if  at  all.  The  anterior  margins 
of  the  disc  are  nearly  straight,  the  posterior  mar- 
gins are  only  slightly  convex,  and  the  posterior 
corners  are  abruptly  rounded  or  even  angular. 
The  tail,  measured  from  the  center  of  the  cloaca, 
is  about  2%  times  as  long  as  the  body  from  cloaca 
to  snout.  The  lower  side  of  the  tail  has  a  narrow 
fold  of  skin  extending  rearward  from  below  the 
origin  of  the  tail  spine  for  a  distance  about  as 

80  This  ray  was  mentioned  as  Dasybatus  marinus  and  as  D.  hasiatus  in  the 
first  edition  of  this  book.  But  the  specimens  in  question  all  belong  to  one 
species,  the  correct  scientific  name  for  which  is  Dasyatis  centroura,  proposed 
by  Mitchill  in  1815,  as  Raja  centroura. 


FISHES   OF  THE  GULF  OF   MAINE 


75 


Fionas  32. — Sting  ray  (Dasyatis  centroura),  male,  about  55^  inches  wide,  Woods  Hole,  Massachusetts,  and  tubercle  from 
tail,  about  0.7  times  natural  size.     From  Bigelow  and  Schroeder.     Drawings  by  E.  N.  Fischer. 


long  as  from  its  own  origin  to  the  cloaca;  the 
upper  side  of  the  tail  is  rounded,  except  for  a  low 
ridge  close  behind  the  spine  (or  spines).  The 
spear-pointed  tail  spines,  of  which  there  are  1,  2, 
or  sometimes  3,  are  situated  well  back  on  the  tail. 
One  spine  that  we  examined  had  about  40  saw 
teeth  on  each  edge. 

Young  ones  are  smoo  th  skinned  (except  for  the  tail 
spines) .  Larger  ones  develop  1  to  3  irregular  rows 
of  conical  tubercles  along  the  midline  of  the  disc, 
with  others  on  the  shoulders  as  well  as  on  the 
outer  posterior  part  of  the  disc,  and  the  tail  be- 
comes very  rough  finally,  with  conspicuous  thorns 


along  its  whole  length  on  its  upper  side,  and 
rearward  from  abreast  of  the  tail  spines  on  its 
lower  side.     The  lower  side  of  the  disc  is  smooth. 

Large  specimens  are  easily  distinguishable  from 
other  sting  rays  of  our  Atlantic  coast  by  their  very 
thorny  tails  and  by  the  large  tubercles  on  the 
outer  parts  of  their  discs.  Small  ones  on  which 
the  thorns  and  tubercles  have  not  yet  developed, 
are  recognizable  by  the  shape  of  the  disc,  com- 
bined with  the  presence  of  a  skin  fold  on  the  lower 
side  of  the  tail  but  none  on  the  upper  side. 

Size. — This  appears  to  be  the  largest  sting  ray 
of   the   western   North   Atlantic.     The   greatest 


76 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


measured  width  definitely  recorded  for  it  is  5  feet, 
the  greatest  measured  length  10  feet  3  inches. 
But  some  certainly  grow  considerably  larger,  for 
a  New  Jersey  specimen  has  been  reported  as 
nearly  7  feet  across;  the  corresponding  length 
would  be  13  to  14  feet,  if  the  tail  were  intact. 

Color. — Fresh  caught  specimens  seen  by  us  at 
Woods  Hole  have  been  dark  brownish  above 
with  the  tail  black  from  the  spine  rearward; 
white  below. 

General  range. — Coastal  waters  of  the  western 
Atlantic,  from  the  latitude  of  Cape  Hatteras 
to  Cape  Cod;  most  common  from  Delaware  Bay 
to  the  Woods  Hole  region. 

Occurrence  in  the   Gulf  of  Maine. — The   only 


claim  of  this  sting  ray  to  mention  here  is  that  one 
was  reported  from  Chatham  on  the  outer  shore  of 
Cape  Cod  many  years  ago,  and  that  it  is  said  to 
have  been  seen  on  the  shoaler  parts  of  Georges 
Bank.  It  has  no  real  status  as  a  Gulf  of  Maine 
fish,  appearing  there  only  as  a  summer  straggler 
from  the  south,  though  it  is  so  common  near 
Woods  Hole  that  the  fish  traps  there  catch  some 
400  to  500  of  them  in  ordinary  summers.81 

Beware  of  handling  any  skate-like  fish  with  a 
long,  whip-like  tail,  lest  it  prove  to  be  a  sting  ray. 
The  tail  spine  (brought  into  action  as  the  tail  i9 
lashed  to  and  fro)  is  a  dangerous  weapon ;  and  the 
wounds  made  by  it  cause  excruciating  pain. 


THE  COW-NOSED  RAYS.     FAMILY  RHINOPTERIDAE 


The  cow-nosed  rays,  like  the  whip-tailed  rays, 
have  a  very  loDg  tail  armed  with  one  or  more 
poisonous  sa wedged  spines;  a  very  flat  broad  disc; 
and  pelvic  fins  with  convex  outer  margins.  But 
their  pectoral  fins  are  interrupted  on  each  side  of 
the  head,  so  that  the  forward  portions  form  a 
separate  two-lobed  fin  extending  forward  from  the 
lower  side  in  front  of  the  mouth  and  nostrils;  their 
crowns  are  high-domed;  their  eyes  and  spiracles 
are  on  the  sides  of  the  head  instead  of  on  its  upper 
surface;  and  they  have  a  small  dorsal  fin  on  the 
upper  side  of  the  tail  in  front  of  the  tail  spines. 
Their  teeth  have  the  form  of  large,  flat  grinding 
plates,  fitting  close  together  in  mosaic  arrange- 
ment; and  there  are  only  7  to  9  series  of  them  in 
each  jaw. 

Cow-nosed  ray 

Rhinoptera  bonasus  (Mitchill)  1815 

Bigelow  and  Schroeder,  1953,  p.  469. 

Garman,  1913,  pi.  37,  as  Rhinoptera  quadriloba. 

Description. — The  cow-nosed  rays  with  all  their 
close  relatives82  have  such  characteristic  out- 
lines, the  shape  of  their  heads  is  so  peculiar  with 
the  eyes  and  spiracles  on  the  sides,  and  their  large, 
flat,  plate-like  teeth  are  so  different  from  those  of 
any  other  Gulf  of  Maine  skates  or  rays  that  they 
are  not  apt  to  be  mistaken  for  anything    else. 

"  This  estimate  is  based  on  our  own  observations  near  Woods  Hole. 

11  The  eagle  rays,  family  Myliobatidae,  and  butterfly  rays,  family 
Oymnurldae,  are  close  allies  of  the  cow-nosed  rays;  none  of  them  has  yet 
been  encountered  in  our  Gulf. 


The  species  in  question  is  characterized  among  its 
confreres  by  the  indented  contour  of  the  front  of 
its  head,  and  by  the  conspicuously  bilobed  outline 
of  the  short  so-called  "subrostral"  fin  that  pro- 
jects forward  from  the  lower  side  of  the  latter. 
The  outer  corners  of  the  pectorals  are  pointed,  and 
their  posterior  margins  distinctly  concave.  The 
pelvic  fins  are  small,  reaching  but  a  short  distance 
back  of  the  posterior  corners  of  the  pectorals. 
The  dorsal  fin  is  rounded  above,  originating  about 
opposite  the  rear  ends  of  the  bases  of  the  pelvics. 
The  tail  measured  from  the  center  of  the  cloaca  is 
about  twice  as  long  as  the  body  from  cloaca  to- 
front  of  head  on  adults  if  not  damaged,  nearly  3 
times  on  small  specimens.  The  tail  spines  (1  or  2) 
are  close  behind  the  rear  limits  of  the  pelvic  fins, 
and  thus  much  further  forward  on  the  tail  than 
those  of  the  sting  rays  (p.  74).  There  usually  are 
7  series  of  teeth  in  each  jaw,  with  up  to  11  to  13 
rows  exposed,  and  in  function  simultaneously. 

Size. — The  cow-nosed  ray  has  been  said  to 
grow  to  a  breadth  of  7  feet.  But  the  largest 
specimen  the  width  of  which  has  either  been 
actually  measured  or  can  be  calculated  from  some 
other  dimension,  was  only  about  38  inches  wide.89 

Color.— Brownish  above,  white  or  yellowish 
white  below.  Some  of  them  are  marked  both 
above  and  below  with  many  narrow  faint  dark 
lines  radiating  out  from  the  center  of  the  disc. 

General  range. — -Western  Atlantic  coast  from 
middle  Brazil  to  southern  New  England. 

<*  Calculated  from  the  dimensions  of  the  head  of  one  from  Rio  de  Janeiro. 


FISHES  OF  THE  GULF  OF   MAINE 


77 


Figure  33. — Cow-nosed  ray  (Rhinoptera  bonasus),  about  22  inches  wide,  Newport,  Rhode  Island. 

Schroeder.     Drawing  by  E.  N.  Fischer. 


From  Bigelow  and 


Occurrence  in  the  Gulf  of  Maine. — The  cow-nosed 
ray  has  even  less  claim  than  the  sting  ray  to  be 
called  a  Gulf  of  Maine  fish,  for  while  schools  of 
them  appear  occasionally  near  Woods  Hole  where 


145  of  them  were  taken  in  the  fish  traps  in  one 
day  on  one  occasion,  and  while  it  is  recorded  from 
Nantucket,  it  has  never  been  seen,  actually,  east 
or  north  of  the  elbow  of  Cape  Cod. 


DEVIL  RAYS.     FAMILY  MOBULIDAE 


The  devil  rays,  like  the  sting  rays  (p.  74)  and 
cow-nosed  rays  (p.  76)  have  the  pectoral  fins 
interrupted  along  the  sides  of  the  head  close  behind 
the  eyes.  But  they  differ  very  noticeably  from 
the  others  mentioned  above  in  the  shape  of  the 
anterior  parts  of  the  pectorals,  for  these  are  in  the 
form  of  two  separate  narrow  ear-like  fins,  set 
vertically  and  curving  forward  from  the  front  of 
the  head.  They  are  further  unique  among  skates 
and  rays  in  the  fact  that  they  feed  on  small 
pelagic  animals,  which  they  sift,  by  a  complex 
sieve-like  modification  of  their  gill  arches,  out  of 
the  water  that  is  gulped  in  by  the  mouth  and 
passed  out  via  the  gill  clefts.  Some  of  them  are 
the  largest  of  the  rays  and  among  the  largest  of 


fishes.  Being  tropical-subtropical  in  nature  they 
have  no  real  place  in  the  fish  fauna  of  our  Gulf, 
but  Manta,  the  largest  of  them  all,  has  been  known 
to  reach  Georges  Bank  as  a  stray  from  warmer 
latitudes. 

Devil  ray  Manta  birostris  (Donndorff)   1798 

Bigelow  and  Schroeder,  1953,  p.  502. 

Description.- — The  so-called  cephahc  fins  of  the 
devil  ray,  pointing  forward,  give  it  so  distinctive 
an  appearance  that  it  could  not  be  confused  with 
any  other  fish,  except  for  some  other  member  of  its 
own  family.  And  it  is  marked  off  from  all  others 
of  these  that  are  known  in  the  Atlantic  by  the 


78 


FISHERY   BULLETIN    OF   THE    FISH   AND  WILDLIFE   SERVICE 


it.      WM 


Figube  34. — Devil  ray  (Mania  birostris),  juvenile  male,  11  feet  5H  inches  wide,  Bimini,  Bahamas. 

Schroeder.     Drawing  by  E.  N.  Fischer. 


From  Bigelow  and 


position  of  its  mouth,  which  extends  across  the 
front  of  its  head  instead  of  being  on  the  under  side. 
Its  cephalic  fins  are  about  one-half  as  broad  at 
the  base  as  they  are  long,  with  thin  lower  edges 
and  thick  fleshy  upper  edges  and  rounded  tips,  and 
each  arises  nearly  vertical  from  the  side  of  the 
head.  When  the  owner  is  swimming  they  point 
directly  forward,  but  when  the  ray  is  feeding,  they 
can  be  curved  inward,  one  toward  the  other 
until  their  tips  nearly  meet  in  front  of  the  mouth. 
The  disc  (not  counting  the  cephalic  fins)  is  a  little 
more  than  twice  as  broad  as  long,  with  tapering 
outer  corners.  The  tail  measured  from  the 
cloaca  is  at  least  as  long  as  the  body  from  cloaca 
to  front  of  head  and  perhaps  longer  still  if  not 
damaged.  And  it  bears  a  small  rounded  dorsal  fin 
on  its  base.  Some  specimens  have  been  described 
as  having  one  or  two  small  tail  spines  close 
behind  the  dorsal  fins.  However,  those  that  we 
have  seen  have  had  none,  but  a  rounded  knob  in  its 
place,  supported  by  a  mass  of  bony  tissue  with  a 
minute  pointed  spur  on  the  upper  side  that  can 
be  felt  but  does  not  break  the  skin.  The  skin  of 
disc,  pelvic  fin,  and  tail  is  roughened  with  small 
tubercles,  below  as  well  as  above.    The  mouth  is 


very  wide,  extending  across  a  little  more  than 
one-half  the  whole  breadth  of  the  front  of  the  head. 
And  the  teeth,  the  lower  jaw  only,  are  minute  and 
very  numerous;  we  counted  about  270  series  in 
about  12  to  18  rows  or  a  total  of  about  4,500  in 
one  specimen  about  11%  feet  wide.  The  gill 
openings  are  noticeably  long. 

Color. — The  upper  side  varies  from  reddish  or 
olive  brown  to  bluish  slate  colored  or  black,  either 
plain  or  with  various  white  markings.  The  lower 
side  is  white  toward  the  center  of  the  disc  but 
gray  around  the  margins,  and  there  may  be 
various  dark  blotches  in  the  region  of  the  gills 
and  on  the  abdomen.  The  rear  part  of  the  tail 
is  gray. 

Size. — This  giant  ray  matures  when  about  14 
to  15  feet  wide.  They  commonly  grow  to  18  feet 
or  so,  and  there  are  recent  records  of  measured 
specimens  19  feet  8  inches,  21  feet  2  inches,  and 
22  feet  wide.  One  14  feet  wide  weighed  1,686 
pounds,  one  from  the  Galapagos  Islands,  18  feet 
wide,  2,310  pounds;  and  one  of  20  feet  taken  long 
ago  off  Venezuela  weighed  3,502  pounds. 

General  range. — Manta  is  known  in  the  Atlantic 
from  middle  Brazil  to  the  Carolinas  and  as  a  rare 


FISHES   OF  THE  GULF  OF   MAINE 


79 


stray  to  southern  New  England  and  Georges  Bank; 
from  Bermuda;  from  Madeira;  and  from  tropical 
West  Africa.  Mantas  are  also  widespread  in  the 
tropical-subtropical  belt  of  the  Pacific  and  Indian 
Oceans,  but  it  is  not  yet  known  whether  they  are 
identical  with  the  Atlantic  species  or  not. 

Occurrence  in  the  Gulf  of  Maine. — The  only 
reason  for  mentioning  this  giant  ray  here  is  that 
a  pair,  judged  to  be   18  to   19  feet  wide,  were 


encountered  on  the  southeast  part  of  Georges 
Bank  late  in  August  1949,  by  Capt.  Henry  W. 
Klimm,  while  out  after  swordfish,  and  so  close  at 
hand  that  their  cephalic  fins  and  purplish  color 
were  noted.  The  nearest  record  to  the  westward 
and  southward  is  of  one  19  feet  wide,  weighing 
1,686  pounds,  harpooned  by  a  sword  fisherman 
a  few  miles  off  Block  Island  and  landed  there  in 
August  192 1.84 


Chimaeroids.    Subclass  Holocephali 


The  chimaeroids,  being  cartilaginous  fishes,  are 
allied  to  the  sharks,  skates  and  rays,  but  are 
separated  from  them  by  many  important  ana- 
tomic characters.  Most  obvious  of  these  externally 
are  that  they  have  no  spiracle;  that  they  have 
only  one  external  gill  opening  on  either  side; 
that  their  tails  are  symmetrical;  and  that  their 
gill  filaments  are  free  at  the  tips  like  those  of  bony 
fishes.  The  chimaeroids  remotely  suggest  the 
grenadiers  in  general  body  form  (p.  243),  but  are 
easily  separable  from  them  at  a  glance;  first  of  all 
by  the  softness  of  their  bodies  and  by  their  naked 


skins,  also  by  the  location  of  the  pelvic  fins  which 
are  set  far  back  under  or  behind  the  tips  of  the 
pectorals,  and  by  the  large  size  of  the  pectoral 
fins,  to  list  only  the  most  obvious  differences. 
There  is  no  danger  of  confusing  them  with  any 
other  Gulf  of  Maine  fishes,  so  curious  is  their 
appearance. 

They  lay  eggs  that  are  astonishingly  large  for 
the  size  of  the  parent  fish,  and  enclosed  in  brown 
horny  capsules  which  are  elliptical,  spindle-shaped 
or  tadpole-shaped  in  different  species.  But 
fertilization  is  internal. 


The  Chimaeras.     Order  Chimaerae 
FAMILY  CHIMAERIDAE 


Chimaera  Hydrolagus  affinis  (Brito  Capello) 
1868 

Bigelow  and  Schroeder,  1953,  p.  539 

Description. — This  species  of  chimaeroid,  the 
only  one  known  from  within  the  geographic  limits 
of  the  Gulf,  is  deepest  (one-sixth  to  one-seventh 
as  deep  as  long)  just  behind  the  gills,  tapers 
gradually  backward  to  a  weak  slender  tail,  and  is 
very  soft-bodied.     The  head  is  short,  its  dorsal 


profile  oblique,  the  snout  conical  with  a  blunt  tip. 
The  forehead  of  the  male  bears  a  curious  cartilag- 
inous hook,  armed  with  recurved  prickles  on  its 
lower  surface,  which  probably  serves  to  clasp  the 
female.  The  mouth,  on  the  lower  side  of  the 
head,  is  small,  with  thick  fleshy  lips;  the  upper 
jaw  is  armed  with  4  flat  plates  in  place  of  teeth, 

«  Reported  by  Gudger  (Science,  N.  Ser.,  vol.  55,  1922,  p.  339).  There  are 
photographs  of  this  specimen  in  the  American  Museum  of  Natural  History 
in  New  York. 


Figure  35. — Chimaera  (Hydrolagus  affinis),  female,  about  31J4  inches  long,  Banquereau  Bank.     From  Bigelow  and 

Schroeder.     Drawing  by  E.  N.  Fischer. 


80 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


the  marginal  pair  set  edgewise,  the  lower  jaw  with 
a  pair  of  marginal  plates  set  edgewise.  The  gill 
openings  are  vertical,  set  very  low  down  on  the 
sides  of  the  neck,  and  each  is  covered  with  a  flap 
of  skin,  paralleling  the  gill  cover  of  bony  fishes. 

There  are  two  distinct  dorsal  fins.  The  first  of 
these  originates  about  over  the  gill  openings,  is 
triangular,  about  as  high  as  long,  and  supported 
at  its  anterior  margin  by  a  stout  spine  that  is  free 
along  the  terminal  part,  with  the  rear  surface  of 
the  free  part  double  saw-edged.  The  second 
dorsal  is  separated  from  the  first  by  a  space  vari- 
able in  length,  and  is  less  than  one-third  as  high 
as  the  first,  with  straight  margin.  The  small 
caudal  fin,  marked  off  from  the  second  dorsal  by 
a  deep  notch,  is  lanceolate  in  shape,  ending  as  a 
short,  whiplike  filament;  and  it  extends  a  short 
distance  forward  along  the  ventral  surface  of  the 
trunk,  there  being  no  separate  anal  fin.  The 
pelvics  and  pectorals  both  have  pointed  tips,  the 
latter  being  much  the  larger  and  reaching  back 
nearly  to  the  point  of  origin  of  the  pelvics.  The 
male  has  a  trifid  copulatory  organ  arising  from  the 
base  of  each  pelvic  fin  on  the  inner  side,  and  also 
a  supplementary  bladelike  clasping  organ  close 
in  front  of  each  pelvic  fin,  its  margin  armed  with 
4  or  5  hooks,  and  lying  in  a  pocket  from  which  it 
can  be  protruded.  The  skin  is  smooth;  the  lateral- 
line  system  is  well  developed  and  ramifies  over  the 
head  in  several  branches. 

This  species  is  a  close  ally  of  the  well  known 
chimaera  of  north  European  seas  (Chimaera 
monstrosa),  but  is  distinguishable  from  it  by  the 
fact  that  it  has  no  separate  anal  fin;  that  there 
is  a  considerable  free  space  between  its  two  dorsal 
fins;  that  the  outline  of  the  second  dorsal  fin  is 


straight;  that  its  caudal  filament  is  much  shorter; 
and  that  its  pectorals  hardly  reach  back  to  the 
pelvics. 

Color. — Lead  color,  tan-brown  or  dark  sepia 
below  as  well  as  above,  except  paler  on  the  throat 
and  grayish  on  the  snout.  The  margin  of  the 
first  dorsal,  the  rear  and  inner  margins  of  the 
pelvics,  and  the  rear  margins  of  the  pectorals  are 
dark. 

Size. — The  largest  specimen  yet  reported,  taken 
85  miles  off  Cape  Sable,  Nova  Scotia,  at  a  depth  of 
between  400  and  500  fathoms,  was  49  inches  long 
and  weighed  17%  pounds  dressed. 

General  range. — Not  uncommon  on  the  conti- 
nental slope  of  North  America  from  the  latitude 
of  Cape  Cod  northeastward,  along  the  Nova  Scotia 
Banks,  to  the  Grand  Banks,  in  160  fathoms  to 
more  than  1,200  fathoms;  also  in  the  eastern  side 
of  the  Atlantic  off  the  coast  of  Portugal. 

Occurrence  in  the  Gulf  of  Maine. — Our  only 
reason  for  mentioning  this  chimaera  is  that  it  is 
(or  was)  so  plentiful  along  the  offshore  slopes  of 
the  Banks  off  the  eastern  part  of  the  Gulf  and  off 
Nova  Scotia  that  many  were  brought  in  for  a 
few  years  subsequent  to  1875,  when  fishermen 
long  fining  for  halibut  extended  their  operations 
down  to  300  fathoms  or  so.  Only  one  seems  to 
have  been  reported  during  the  past  25  years, 
caught  off  Browns  Bank,  85  miles  southwest  of 
Cape  Sable,  between  400  and  500  fathoms  on 
October  15,  1930.8S  But  perhaps  it  would  be 
found  no  less  plentiful  now  than  of  old,  if  sought 
at  the  proper  depth.  The  shoalest  capture  of 
which  we  found  record  was  at  160  fathoms. 
Nothing  is  known  of  its  way  of  life  nor  have  its 
egg  cases  been  seen. 


THE  BONY  FISHES.     CLASS  OSTEICHTHYES 
THE  STURGEONS.     FAMILY  ACIPENSERIDAE 


The  sturgeons,  like  the  sharks,  have  an  uneven 
{"heterocercal")  tail  with  the  vertebral  column 
extending  out  along  the  upper  lobe.  But  there  is 
no  danger  of  mistaking  a  sturgeon  for  a  shark  for 
it  has  only  one  gill  opening  on  each  side,  while  the 
gills  are  enclosed  by  bony  gill  covers.  And  the 
combination  of  gills  of  this  kind  with  sharklike 
tail  and  with  the  fact  that  the  head  is  covered  by 


bony  plates  united  by  sutures,  sets  the  sturgeons 
off  from  all  other  Gulf  of  Maine  members  of  their 
own  class.  Two  species  of  sturgeons  are  known 
from  the  Gulf,  one  of  which  once  was  rather 
common  there;  the  other  is  extremely  scarce 
everywhere. 

«  Reported  by  Firth,  Bull.  Boston  Soc.  Nat.  Hist.,  61, 1931,  p.  9.   It  was  49 
inches  long  and  weighed  17H  pounds  dressed. 


FISHES   OF  THE  GULF  OF   MAINE 


81 


2. 


KEY  TO  GULF  OF  MAINE  STURGEONS 

The  successive  bucklers  in  the  dorsal  row  touch  each  other  or  even  overlap;  the  space  between  the  dorsal  row  of  buck- 
lers and  the  uppermost  of  the  two  lateral  rows  is  thickly  set  with  coarse  prickles Sea  sturgeon,  p.  81. 

The  successive  bucklers  in  the  dorsal  row  are  separated  one  from  the  next  by  spaces  up  to  %  as  long  as  the  bucklers; 
the  space  between  the  dorsal  row  of  bucklers  and  the  uppermost  of  the  two  lateral  rows  is  only  sparsely  strewn 
with  fine  prickles Short-nosed  sturgeon,  p.  84. 


Sea  sturgeon  Acipenser  sturio  Linnaeus,  1758  86 
Jordan  and  Evermann,  1896-1900,  p.  105 

Description. — The  skin  of  the  sturgeon  is 
armored  with  a  row  of  large  bony  shields  or 
bucklers  along  the  middle  of  its  back  (the  succes- 
sive bucklers  touching  or  even  overlapping)  with 
a  second  row  of  smaller  bucklers  high  up  along 
each  side  of  the  body;  and  with  a  third  row,  also 
smaller,  lower  down,  along  the  line  of  transition 
from  side  to  belly.  Each  buckler  has  a  longitu- 
dinal keel  with  a  spur,  which  is  so  sharp  on  small 
fish  that  these  are  hard  to  handle,  lower  and 
blunter  on  large.  On  the  average  there  are  10 
or  11  (10 — 16)  bucklers  in  the  mid-dorsal  row; 
28  or  29  (26 — 34)  in  each  upper  lateral  row;  and 
9  to  14  in  each  of  the  lower  lateral  rows.  The 
dorsal  row  runs  from  above  the  gill  covers  back 
to  the  dorsal  fin,  and  each  of  the  dorsal  shields 
reaches  to  the  next  shield  or  even  overlaps  it. 
The  upper  lateral  rows  run  from  the  gill  openings 
back  to  the  root  of  the  tail  fin;  the  lower  lateral 
rows  from  close  behind  the  pectoral  fin  to  the 
pelvic  fin,  also  from  the  pelvic  fin  back  as  far  as 
the  anal  fin.  And  each  shield  in  each  of  the  two 
lateral  rows  is  separated  from  the  next  shield  by 
a  space  up  to  one-half  as  long  as  the  shields. 

The  body  is  rather  slender  and  rendered  more 
or  less  pentagonal  in  cross  section  by  the  five  rows 
of  shields,  instead  of  rounded  as  it  is  in  the  majority 
of  bony  fishes.  The  snout  is  narrow  in  young 
sturgeons  less  than  2  to  2%  feet  long,  depressed 
below  the  level  of  the  forehead,  nearly  flat  below, 

M  It  still  is  an  open  question,  that  we  cannot  answer,  whether  the  sea 
sturgeon  of  eastern  North  America  is  identical  with  the  European  sea  stur- 
geon, Is  a  recognizable  race  of  the  latter,  or  is  a  separate  species;  if  the 
last,  its  scientific  name  is  Acipenser  oxyrinchus  Mitchill,  1816. 


and  longer  (from  the  eyes  forward)  than  the  dis- 
tance is  from  the  eyes  rearward  to  the  upper 
corners  of  the  gill  openings.  But  it  changes 
shape  as  the  fish  grows,  becoming  blunter,  straight 
in  dorsal  profile,  and  considerably  shorter  rela- 
tively. The  mouth,  situated  on  the  under  side 
of  the  head,  is  small,  toothless  (except  in  larval 
stages),  with  protractile  lobed  lips,  and  there  are 
four  pointed  barbels  in  a  row  across  the  lower 
side  of  the  snout  in  front  of  the  mouth.  The 
single  rather  small  triangular  dorsal  fin  stands 
far  back,  with  its  rear  edge  over  that  of  the  still 
smaller  anal  fin.  The  ventral  fins  are  likewise  far 
back.  The  pectorals  are  set  almost  as  low  as  the 
plane  of  the  belly.87 

Color. — Olive  greenish  or  bluish  gray  above, 
gradually  fading  on  the  sides  and  changing 
rather  abruptly  below  the  upper  lateral  rows  of 
shields  to  the  white  of  the  belly. 

Size. — The  sea  sturgeon  is  a  very  large  fish. 
In  the  Delaware  River  where  sturgeon  persisted 
until  recently  in  larger  numbers  than  in  New 
England,  ripe  males  are  up  to  about  6  to  7  feet 
in  length,  averaging  65  pounds  in  weight;  the 
spawning  females  (which  are  larger),  up  to  about 
10  feet  and  to  about  250  pounds,88  with  a  larger 
one  taken  from  time  to  time.  And  the  general 
run  was  about  the  same  in  the  Kennebec,  to  judge 
from  an  average  weight  of  120  pounds  for  males 
and  females  together,  during  the  years  when  a 
fishery  was  carried  on  there.  But  some  still  grow 
considerably  larger  in  Gulf  of  Maine  waters. 
Thus  9  weighing  between  350  pounds  and  600 

«'  Vladykov  and  Beaulieu  (Natural.  Canad.,  vol.  73,  1946,  pp.  143-204), 
give  a  detailed  account  of  the  characters  that  separate  the  sea  sturgeon  from 
the  lake  sturgeon  (Acipenter  fulvescens  Raflnesque,  1817). 

»  According  to  Cobb,  Rept.  U.  S.  Fish  Comm.  (1899),  1900,  p.  277. 


*fr~^B^3^  ^si1-^*!* 


Figure  36. — Sturgeon  {Acipenser  sturio),  Potomac  River  specimen.     From  Goode.     Drawing  by  H.  L.  Todd. 


82 


FISHERY   BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


pounds  were  landed  in  Portland,  Maine,  from  the 
South  Channel,  Georges  Bank,  Browns  Bank, 
and  Western  Bank  off  Nova  Scotia  during  the 
period  1927-1935.89  About  12  feet  is  perhaps 
the  greatest  length  to  be  expected  today.  But 
18  feet,  reported  for  New  England  many  years 
ago,  may  not  have  been  an  exaggeration,  for 
sturgeon  as  long  as  that  have  been  reported  from 
Europe  also.  The  heaviest  Gulf  of  Maine  stur- 
geon reliably  reported  (to  our  knowledge)  was 
one  of  600  pounds,  landed  in  Portland  by  the 
steam  trawler  Fabia  from  Georges  Bank,  Decem- 
ber 21,  1932. 

The  following  relationship  between  length  and 
weight,  for  sea  sturgeons  up  to  7%  feet  long,  taken 
in  the  lower  St.  Lawrence  River,90  would  probably 
apply  to  Gulf  of  Maine  fish,  equally:  7  to  9  pounds 
at  30  inches  (to  fork  of  tail);  15  to  18  pounds  at 
40  inches;  about  35  pounds  at  50  inches;  55  to  57 
pounds  at  5  feet;  about  100  pounds  at  6  feet;  and 
about  190  pounds  at  7%  feet. 

Habits. — The  sturgeon  makes  most  of  its  growth 
in  salt  water  but  enters  fresh-water  rivers  to  spawn, 
as  do  the  salmon,  the  shad,  and  the  alewife.  The 
large  adult  fish  enter  (or  once  entered)  the  Gulf  of 
Maine  rivers  late  in  the  spring,  working  their  way 
slowly  upstream  beyond  tidewater  before  deposit- 
ing their  eggs.  So  far  as  known,  spawning  takes 
place  in  our  rivers  in  May,  June,  and  perhaps  as 
late  as  July.  It  has  been  suggested  that  some  may 
spawn  in  brackish  water  from  the  fact  that  females 
with  large  eggs  have  been  taken  near  Woods  Hole 
in  June  and  July  (i.  e.,  in  the  spawning  season). 
Spawning  leaves  the  spent  "cows"  in  very  poor 
condition.  In  the  Delaware,  however,  and  pre- 
sumably in  Gulf  of  Maine  rivers,  they  "become 
again  quite  plump,  acquiring  considerable  addi- 
tional weight"  91  before  they  go  down  stream  again, 
which  some  of  them  do  not  do  until  September, 
according  to  observations  in  the  Delaware.  But 
we  do  not  know  how  many  years  in  succession  a 
given  fish  may  spawn. 

A  single  female  may  produce  as  many  as 
2,400,000  eggs  which  hatch  in  about  a  week  after 
they  are  fertilized.92  Judging  from  European 
observations  on  artificially  reared  sea  sturgeon, 

»  Records  collected  by  the  late  Walter  H.  Rich  of  the  U.  S.  Bureau  of 
Fisheries. 

*•  According  to  measurements  and  weights  of  1,592  sturgeons  by  Vladykov, 
Rapp.  Gen.  Minlstr.  Chasse.  PGch.,  Quebec  (1948-1949),  1949,  pp.  43-54. 

•'  Ryder,  Bull.  U.  S.  Fish  Comm.,  vol.  8,  1890,  p.  266. 

«  Ryder  (Bull.  D.  S.  Fish  Comm.,  vol.  8, 1890,  p.  231)  describes  the  spawn- 
ing and  early  development  of  the  sturgeon  in  the  Delaware  River. 


the  larvae  may  be  expected  to  grow  to  12  mm.  in 
length  within  5  days  after  hatching;  to  16-17  mm. 
in  2  weeks;  to  20  mm.  in  4  weeks;  and  to  4-5 % 
inches  in  2  months. 

Some  young  sturgeon  may  live  several  years  in 
the  lower  tidal  reaches  of  the  rivers  in  which  they 
are  spawned,  until  they  have  grown  to  a  length 
of  2)i  to  3  feet,  as  appears  to  be  the  case  in  the 
Hudson.93  And  it  seems  that  they  pass  their 
entire  growth  period  in  the  salt  estuary  of  the  St. 
Lawrence  River,  for  sturgeons  are  taken  there  of 
all  sizes  from  a  few  inches  long  up  to  7-8  feet  or 
longer.94  But  others  may  descend  during  their 
first  year,  for  sperlets  only  5  to  6  inches  long  have 
been  found  at  the  mouth  of  the  Delaware  River 
and  of  the  Elbe  in  Europe.95 

Some  Gulf  of  Maine  sturgeon  have  taken  to  the 
sea  by  the  time  they  have  grown  to  3  feet  or  so,  as 
proved  by  the  capture  of  sturgeons  of  that  size 
at  various  points  around  the  coasts  of  the  Gulf, 
and  off  southern  New  England.  And  recent  ob- 
servations in  the  Hudson  by  Greeley  make  it 
likely  that  all  the  sturgeon  that  are  spawned  in 
rivers  emptying  into  the  Gulf  of  Maine  go  to  sea 
sooner  or  later  to  complete  their  growth.99 

Sturgeon  grow  rather  slowly  at  first  while  still 
in  their  parent  streams.  Four,  for  example, 
that  were  tagged  in  the  lower  St.  Lawrence  when 
29  to  33  inches  long,  and  recaptured  nearby  2 
to  3%  years  later,  had  gained  only  about  2  to  5 
inches  in  length  per  year.97  Very  slow  growth 
is  also  indicated  by  ages  of  5  to  6  years  at  24  to  28 
inches;  7  years  at  25  to  31  inches;  and  8  years  at 
32  to  34  inches,  for  sturgeon  from  the  tidal  waters 
of  the  lower  Hudson,  as  estimated  from  the  mark- 
ings on  their  otoliths.98  It  also  seems  that 
sturgeon,  like  many  other  fish,  make  most  of  their 
growth  during  the  warm  season  in  such  situations 
for  one  marked  fish  in  the  Elbe  did  not  grow  at  all 
between  November  and  the  following  February, 
whereas  a  second  grew  from  17  cm.  (6%  in.)  to  38 
cm.  (15  in.)  in  length  between  January  17  and 

>'  See  Greeley  (Supp.  26  Ann.  Rept.  Conserv.  Dept.  New  York,  1937,  pp. 
68.  78-82,  89)  for  a  study  of  the  sturgeon  in  the  Hudson  River. 

■<  A  series  of  1,592  sea  sturgeons  from  the  lower  St.  Lawrence  River,  studied 
by  Vladykov  (Rapp.  Gen.  Minstr.  Chasse,  P8ch.  Quebec  (1948-1949)  1949, 
pp.  53-56)  included  a  good  representative  of  sizes  from  about  4  inches  up  to 
90  inches. 

M  Prince  reports  a  6-rnch  sturgeon  from  Hudson  Bay  (Rept.  Sixty-seventh 
Meeting,  British  Assoc.  Adv.  Sci.,  Toronto,  1897,  p.  687). 

•«  Greeley,  Suppl.  26  Ann.  Rept.  Conserv.  Dept.  New  York,  1937,  p.  82. 

•»  Vladykov  (Rapp.  Gen.  Minlstr.,  Chasse,  Pech.  Quebec,  1948-1949, 
pp.  61-63,  66,  table  19). 

•»  Greeley,  Supp.  26  Ann.  Rept.  Conserv.  Dept.  New  York,  1937,  p.  68, 
table  10. 


FISHES   OF  THE  GULF  OF   MAINE 


83 


the  following  April,  and  a  third  from  43%  cm. 
(17%  in.)  to  64  cm.  (25%  in.)  from  April  9  to  the 
following  December.  But  sturgeon  grow  much 
more  rapidly  after  they  go  to  sea,  if  ages  (esti- 
mated from  otoliths)  of  11  years  for  a  75-inch 
sturgeon,  and  12  years  for  two  others  of  88  and 
100  inches  are  anywhere  near  the  truth.09 

The  sturgeon  is  a  bottom  feeder,  rooting  in  the 
sand  or  mud  with  its  snout  like  a  pig  (the  barbels 
serving  as  organs  of  touch)  as  it  noses  up  the  worms 
and  mollusks  on  which  it  feeds  and  which  it  sucks 
into  its  toothless  mouth  with  considerable  amounts 
of  mud.  It  also  consumes  small  fishes,  particu- 
larly sand  launce.  Small  ones,  while  living  in 
estuaries  and  around  river  mouths,  subsist  largely 
on  amphipod  and  isopod  Crustacea.  Sturgeon, 
like  salmon,  eat  little  or  nothing  while  traveling 
up  river  to  spawn. 

When  at  ease  sturgeon  swim  slowly  to  and  fro, 
seeming  very  sluggish.  But  they  are  capable 
of  darting  ahead  like  an  arrow  on  occasion,  and 
they  often  come  to  the  surface  to  jump  clear  of 
the  water.  Though  they  usually  offer  no  resist- 
ance when  netted,  large  ones  are  very  strong. 

General  range.— Coastal  waters  from  the  St. 
Lawrence  River  to  the  Gulf  of  Mexico,  running 
up  into  rivers  to  spawn;  reported  from  Hudson 
Bay,  also  Scandinavia  to  the  Mediterranean,  if 
the  American  and  European  sea  sturgeons  belong 
to  the  same  species. 

Occurrence  in  the  Gvlf  of  Maine.— The  sea 
sturgeon  is  (or  was)  well  known  in  the  St.  John, 
Penobscot,  Kennebec,  and  Merrimac  Rivers,  and 
has  even  been  taken  some  distance  from  the 
mouths  of  streams  no  larger  than  the  Charles  River 
and  the  Parker  River  in  Essex  County,  Mass.,1 
where  some  are  still  seen  jumping  in  July  and  one 
is  taken  occasionally.  In  fact,  sturgeon  once 
entered  practically  every  stream  of  any  size 
emptying  into  the  Gulf  of  Maine.  Wood,  writing 
of  Massachusetts  in  1634,"  described  them  as 
"all  over  the  country,  but  best  catching  of  them 
be  upon  the  shoales  of  Cape  Code  and  in  the 
river  of  Merrimacke,  where  much  is  taken, 
pickled  and  brought  for  England,  some  of  these 
be  12,  14  and  18  foote  long."     In  fact,  an  odd 


••  See  footnote  98. 

1  Two  sturgeon  44  and  45M  Inches  long,  netted  in  the  Parker  River  at 
Newbury,  Mass. ,  July  23, 1933,  are  (or  were)  In  the  collection  of  the  Boston 
Boclety  of  Natural  History,  now  the  New  England  Science  Museum  (Bull. 
Boston  Soc.  Nat.  Hist.,  69,  Oct.  1933,  p.  8). 

'»  New  England's  Prospect,  1634,  p.  37. 


sturgeon  still  enters  the  mouth  of  the  Merrimac, 
witness  one  of  230  pounds  netted  there  on  Sep- 
tember 14,  1938  and  landed  in  Newburyport.2 

Sturgeons  may  be  expected  anywhere  off  the 
coasts  of  the  Gulf  of  Maine  during  their  sojourn 
in  salt  water.  There  is  definite  record  of  them  at 
sundry  localities  on  both  sides  of  the  Bay  of  Fundy ; 
off  Mt.  Desert  Island;  in  Penobscot  Bay;  in  Casco 
Bay;  at  the  mouth  of  the  Piscataqua  River;  on  the 
Boars  Head-Isles  of  Shoals  fishing  ground,  where 
several  3  to  4  feet  long  were  taken  in  gill  nets  dur- 
ing April  and  M&j  1913;  at  the  mouths  of  the 
Essex  and  Ipswich  Rivers,  where  jumping  stur- 
geon have  been  reported  recently  in  the  daily 
press ; 8  at  the  mouth  of  Gloucester  Harbor,  where 
an  angler  reports  catching  one  of  about  12  pounds 
while  fishing  for  tautog;  inside  and  outside  Boston 
Harbor;  at  Provincetown ;  off  Truro,  Cape  Cod; 
and  at  Nantucket,  as  well  as  along  the  southern 
New  England  coast  to  the  westward.  Some  also 
extend  their  wanderings  to  the  offshore  fishing 
banks  as  they  grow.  Thirty,  for  example,  rang- 
ing in  weight  from  120  to  600  pounds  were  landed 
in  Portland  and  Boston  by  otter  trawlers  from 
Nantucket  Shoals,  from  South  Channel,  and  from 
Georges  and  Browns  Banks,  during  the  years 
1927-1936.4  Probably  all  of  these  were  on  bot- 
tom when  caught,  to  judge  from  their  diet  (p.  83), 
and  from  the  fact  that  sturgeon  have  been  hooked 
on  cod  and  haddock  lines  as  deep  as  25  fathoms  in 
Scandinavian  waters.  Nothing  beyond  this  is 
known  of  their  movements  in  our  Guff. 

Importance. — It  is  only  the  scarcity  of  the  sea 
sturgeon  in  the  Gulf  of  Maine  that  limits  its  com- 
mercial importance  there  and  in  the  tributary 
rivers.  The  few  taken  are  picked  up  acciden- 
tally in  traps  or  weirs,  in  drift  nets,  or  by  the  otter 
trawlers. 

In  former  years,  when  our  streams  were  less 
obstructed  and  sturgeons  more  plentiful,  the  catch 
was  of  considerable  value  in  some  of  the  larger 
rivers.  It  is  interesting,  for  instance,  to  read  that 
sturgeon,  doubtless  from  the  Kennebec  River  and 
cured  near  what  is  now  Brunswick,  Maine,  were 
shipped  to  Europe  as  early  as  1628;  and  that  large 
quantities  were  also  shipped  to  Europe  from  near 
Ipswich,  Mass.,  in  1635.  In  the  Kennebec,  where 
an  intermittent  fishery  had  long  been  maintained. 

» Reported  In  the  Boston  Globe.  Sept.  15, 1938. 
» The  Boston  Herald,  June  1950. 

<  Reports  collected  by  the  late  Walter  H.  Rich,  U.  S.  Bureau  of  Fisheries, 
and  notices  in  the  dally  press. 


84 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


the  catch  was  about  250  fish  in  1880,  yielding 
12,500  pounds  of  meat,  and  not  much  less  in  1898 
(10,875  pounds).  But  the  yearly  landings  were 
only  about  one  fourth  as  great  there  (2,777  pounds) 
by  1919.  And  the  reported  landings  of  sturgeon 
from  the  entire  coastline  of  Maine  (including  what 
few  were  brought  in  from  offshore)  had  fallen  to 
only  300  pounds  in  1940,  and  400  pounds  in 
1947.  Reported  landings  in  Massachusetts  of 
5,300  pounds  in  1940  (all  by  otter  trawlers)  and 
of  6,600  pounds  (5,000  pounds  by  otter  trawlers, 
from  off  shore),  corresponding  to  some  50  to  70 
fish,  if  they  weighed  as  little  as  100  pounds  each, 
will  further  illustrate  their  present-day  scarcity. 
We  have  never  heard  of  a  large  sturgeon  hooked 
by  an  angler  in  the  Gulf  of  Maine.  But  we  hear 
from  time  to  time  of  a  small  one  caught  in  this 
way,  as  already  remarked  (p.  83)  .5  And  the 
skill  of  a  woman  angler  9  who  foul-hooked  a  stur- 
geon about  6  feet  long,  and  beached  it  on  surf- 
casting  tackle  after  a  long  fight,  fishing  alone  at 
Wasque  Point,  Marthas  Vineyard,  on  July  15, 
1950,  was  widely  heralded  in  the  daily  press. 

Short-nosed  sturgeon  Acipenser  brevirostrum 
LeSueur  1818 

Little  sturgeon 

Jordan  and  Evermann,  1896-1900,  p.  106. 

Description. — The  little  sturgeon  resembles  the 
sea  sturgeon  so  closely  in  general  appearance  that 
we  need  note  only  the  most  obvious  differences. 
These  are  that  the  shields  in  its  dorsal  row  are 
relatively  smaller,  and  that  each  is  separated  from 
the  next  by  a  space  up  to  %  as  long  as  the  shields 
themselves  (successive  dorsal  shields  in  contact  or 
overlapping  in  the  sea  sturgeon);  that  the  space 
between  its  dorsal  row  of  shields  and  the  upper 

•  We  once  saw  one  small  one  about  23  Inches  (575  mm.)  long  foul  hooked  In 
the  side  on"  South  Beach,  New  York,  December  21, 1923,  and  heard  of  a  simi- 
lar experience  by  the  same  angler  a  year  later. 

•  Mrs.  George  T.  Rice.  About  30  others  were  seen  by  her  at  the  same  time 
In  a  deep  slew  formed  by  a  new  bar. 


lateral  row  on  each  side  is  only  sparsely  set  with 
fine  prickles  (closely  set  with  coarse  prickles  in 
the  sea  sturgeon);  and  that  its  viscera  are  black- 
ish (pale  in  the  sea  sturgeon) ;  also  the  number  of 
rays  in  the  anal  fin  averages  smaller  in  the  little 
sturgeon  (19-22)  than  in  the  sea  sturgeon  (23-30). 
The  snout,  too,  is  considerably  shorter  relatively, 
as  well  as  broader,  than  it  is  in  young  sea  stur- 
geons of  equal  size.  And  while  the  snout  is  about 
as  long,  relatively,  in  the  one  species  as  in  the 
other  whea  they  are  full  grown,  sea  sturgeons  are 
then  so  much  the  larger  that  there  is  no  danger  of 
confusing  the  one  kind  with  the  other. 

Color. — Described  as  blackish  above,  tinged 
with  olive  above  the  upper  lateral  line  of  shields, 
marked  with  alternate  black  and  pale  bands;  sides, 
below  the  upper  lateral  row  of  shields,  reddish 
mixed  with  violet;  abdomen  white.7 

Size. —  This  is  a  much  smaller  fish  than  the  sea 
sturgeon.  Males  may  mature  when  only  19-20 
inches  long  and  most  of  them  do  by  the  time  they 
pass  21  inches;  most  of  the  females  at  about  24 
inches.  The  largest  so  far  recorded  is  one  of  about 
36  inches,  in  the  Museum  of  Comparative  Zoology. 
One  about  31  inches  long  weighed  7  pounds  4 
ounces.8 

Habits. — Nothing  is  known  of  the  habits  of  the 
little  sturgeon  except  that  it  spawns  in  rivers  and 
that  it  does  so  late  in  April  in  the  lower  Hudson. 
The  fact  that  fair  sized  specimens  are  taken  there 
in  summer  and  also  in  winter,  suggests  that  it 
may  not  be  as  regularly  migratory  as  the  sea 
sturgeon  is.9  But  the  places  of  capture  of  the 
Gulf  of  Maine  specimens  mentioned  below  show 
that  some  certainly  go  out  into  the  open  sea  and 
wander  for  some  distance  from  their  parent  stream. 

General  range. — So  far  as  we  know,  the  only 


We  have  not  seen  a  fresh-caught  specimen. 

■  For  further  details  as  to  size,  see  Ryder,  Bull.  U.  S.  Fish  Comm.,  vol.8, 
1890,  p.  238;  and  Greeley,  Suppl.  26  Ann.  Kept.  Conserv.  Dept.  New  York, 
1937,  p.  69,  table  11,  pp.  82,  90. 

•  Greeley,  Suppl.  to  26  Ann.  Rept.  Conserv.  Dept.  New  York,  1937,  p.  90, 
makes  this  suggestion. 


Figure  37. — Short-nosed  sturgeon   (Acipenser  brevirostrum),  Woods  Hole  specimen.     From  Goode.     Drawing  from  a 

photograph. 


FISHES   OF  THE  GULF  OF   MAINE 


85 


locality  records  definitely  belonging  to  this  species, 
not  to  young  sea  sturgeons,  are  from  Province- 
town  and  Waquoit,  Mass.;  from  the  Hudson 
River,  N.  Y. ;  from  Delaware  Bay  and  River; 
and  from  Charleston,  S.  C. 

Occurrence  in  the  Gulf  of  Maine. — The  only 
recent  record  of  the  little  sturgeon  in  the  Gulf  is 
of  one  about  23  inches  long,  taken  at  Province- 


town  about  1 907  and  now  mounted  in  the  Museum 
of  Comparative  Zoology.10  The  Museum  of  the 
Essex  Institute,  Salem,  also  has — or  had — a 
stuffed  sturgeon  from  Rockport,  Mass.,  identified 
as  this  species  by  Goode  and  Bean.11  Evidently 
the  sturgeon  is  now  very  scarce  in  our  Gulf  and 
there  is  no  reason  to  think  that  it  ever  has  been 
more  plentiful  there. 


The  Herring  and  Tarpon  Tribes 
FAMILIES  CLUPEIDAE,  DUSSUMIERIIDAE,  AND  ELOPIDAE 


The  true  herrings  (Clupeidae)  are  soft-finned 
fishes  wholly  lacking  spines,  with  one  short  dorsal 
fin,  deeply  forked  tails,  ventral  fins  situated  on 
the  abdomen  far  behind  the  pectorals,  teeth  small 
or  lacking  in  adults,  deep  bodies  flattened  side- 
wise,  and  large  scales  that  slip  off  at  a  touch. 
They  are,  perhaps,  the  most  familiar  of  northern 
sea  fishes  and  certainly  are  the  most  abundant  in 
number  of  individuals.  Seven  species  of  herring 
occur  in  the  Gulf  of  Maine — the  hickory  shad  (not 
very  common),  the  sea  herring,  the  alewife,  the 
blueback,  and  the  shad  (regular  and  plentiful), 
thread  herring  (scarce),  and  the  menhaden  (irreg- 
ular in  its  occurrence) .  The  shad,  menhaden,  sea 
herring,  and  thread  herring  are  easily  named ;  but 
the  alewife  and  the  blueback  resemble  one  another 


so  closely  that  they  are  often  confused,  even  by  the 
fishermen  who  handle  them  constantly.  The  round 
herrings  (Family  Dussumieriidae)  differ  from  the 
true  herrings  chiefly  in  tbeir  rounded  bellies  and 
less  deep  bodies.  The  members  of  the  Tarpon 
Tribe  (Family  Elopidae)  are  very  closely  allied 
to  the  true  herrings  (Clupeidae),  from  which  they 
differ  in  having  a  bony  plate  on  the  throat  between 
the  branches  of  the  lower  jaw.  There  are  only 
about  five  species,  all  of  them  tropical.  Two 
are  known  from  the  Gulf,  as  strays. 


10  This  Museum  also  has  another  of  about  36  Inches  from  Waquoit,  on  the 
southern  shore  of  Massachusetts. 

"  Bull.  Essex  Inst.,  vol.  11,  1879,  p.  27.  A  sturgeon  was  reported  as  brt- 
virostris  from  Boston  Harbor  many  years  ago,  but  there  is  no  way  now  of 
checking  the  Identification. 


KEY  TO  GULF  OF  MAINE  HERRINGS  AND  TARPONS 

1.  Last  dorsal  fin  ray  prolonged 2 

Last  dorsal  ray  not  prolonged 3 

2.  Dorsal  fin  originates  in  advance  of  the  ventrals;  scales  only  moderately  large Thread  herring,  p.  112 

Dorsal  fin  originates  behind  the  ventrals;  scales  very  large Tarpon,  p.  87 

3.  Belly  rounded 4 

Belly  sharp  edged 5 

4.  Scales  very  small;  mouth  very  large  with  upper  jaw-bone  extending  considerably  beyond  the  rear  edge  of  the  eye; 

point  of  origin  of  dorsal  fin  about  over  that  of  the  ventral  fins Ten  pouDder;  p.  86 

Scales  large;  mouth  small,  with  upper  jaw-bone  extending  rearward  only  about  as  far  as  the  front  edge  of  the  eye; 
point  of  origin  of  dorsal  fin  well  in  advance  of  that  of  the  ventral  fins Round  Herring,  p.  87 

5.  Head  (tip  of  snout  to  edge  of  gill  cover)  very  large,  occupying  about  one-third  the  total  length  of  the  body  to  base 

of  the  central  rays  of  the  caudal  fin;  free  edges  of  scales  fluted,  not  rounded Menhaden,  p.  113 

Head  about  one-fourth  the  total  length  of  the  body;  free  edges  of  the  scales  rounded 6 

6.  Distance  from  point  of  origin  of  dorsal  fin  to  tip  of  lower  jaw  (mouth  closed)  about  as  long  as  from  origin  of  dorsal 

fin  to  base  of  central  rays  of  caudal  fin;  edge  of  belly  hardly  saw-toothed,  though  sharp;  general  form  comparatively 

shallow;  there  is  a  cluster  of  teeth  on  the  roof  of  the  mouth Sea  herring,  p.  88 

Distance  from  point  of  origin  of  dorsal  fin  to  tip  of  lower  jaw  (mouth  closed)  considerably  shorter  than  from  point  of 
origin  of  dorsal  fin  to  origin  of  central  rays  of  caudal  fin;  edge  of  belly  more  or  less  strongly  saw-toothed,  especially 
in  space  between  the  ventral  and  anal  fins;  general  form  deep;  there  are  no  teeth  on  the  roof  of  the  mouth.  _       7 

7.  The  tip  of  the  lower  jaw  extends  noticeably  beyond  the  upper  when  mouth  is  closed Hickory  shad,  p.  100 

The  tip  of  the  jaw  does  not  extend  appreciably  beyond  the  upper  when  mouth  is  closed 8 


86 


FISHERY   BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


9. 


The  upper  outline  of  the  forward  part  of  the  lower  jaw  (visible  if  mouth  is  opened)  is  nearly  straight,  ana  does  not  show 

a  pronounced  angle;  the  upper  jaw  extends  back  about  level  with  the  rear  edge  of  the  eye Shad,  p.  108 

The  upper  outline  of  the  forward  part  of  the  lower  jaw  is  concave  with  a  pronounced  angle;  the  upper  jaw  reaches 

back  only  about  to  the  level  of  the  center  of  the  eye 9 

Breadth  of  eye  is  greater  than  distance  from  front  of  eye  to  tip  of  snout;  back  distinctly  grey  green;  lining   of   belly 

cavity  pale  grey Alewife,  p.  101 

Breadth  of  eye  is  only  about  as  great  as  distance  from  front  of  eye  to  tip  of  snout;  back  distinctly  blue  green;  lining 

of  belly  cavity  sooty  or  black Blue  back,  p.  106 


Ten  pounder  Elops  saurus  Linnaeus  1766 
Jordan  and  Evermann,  1896-1900,  p.  410. 

Description. — The  ten  pounder  is  herring-like  in 
the  arrangement  of  its  fins,  with  the  single  and 
soft-rayed  dorsal  fin  originating  about  midway 
along  its  back;  in  having  no  adipose  fin;  in  the 
position  of  its  ventral  fins  about  midway  between 
tip  of  snout  and  fork  of  tail ;  and  in  its  forked-tail 
fin.  But  its  scales  are  very  much  smaller  relatively 
than  those  of  any  of  our  herrings,  and  its  mouth 
is  much  larger,  with  the  upper  jawbone  extending 
rearward  considerably  beyond  the  rear  edge  of  the 
eye.  Being  about  one-sixth  as  deep  as  it  is 
long,  it  is  a  much  more  slender  fish  than  any  of 
our  herrings  except  the  round  herring,  and  its 
belly  is  rounded  like  that  of  the  latter.  But  its 
trunk  is  more  flattened  sidewise  than  that  of  the 
round  herring,  its  dorsal  fin-origin  is  over  the  ven- 
trals  (well  in  advance  of  the  ventrals  in  the  round 
herring),  and  its  tail  fin  is  much  wider  relatively 
than  that  of  any  herring,  and  more  deeply  forked. 

A  more  important  structural  character  is  that 
its  throat  is  stiffened  between  the  branches  of  its 
lower  jaw  by  a  long  bony  plate,  which  it  shares 
with  the  tarpon,  but  which  no  member  of  the  her- 
ring tribe  has.  Its  closest  affinity  among  fishes 
yet  known  from  our  Gulf  is  with  the  tarpon.  But 
its  scales  are  very  much  smaller  than  those  of  the 


latter,  nor  does  its  dorsal  fin  have  the  prolonged 
ray  characteristic  of  the  tarpon. 

Color. — Silvery  all  over,  with  the  back  bluish, 
the  lower  parts  of  the  sides  and  the  lower  surface 
yellowish;  the  dorsal  and  caudal  fins  dusky  yellow- 
ish and  silvery;  the  ventral  and  pectoral  fins 
yellowish  speckled  and  dusky. 

Size. — The  ten  pounder  is  said  to  grow  to  a 
length  of  3  feet,12  but  few  of  those  caught  are 
longer  than  about  20  inches. 

General  range. — Atlantic  coast  of  America,  from 
Brazil  northward;  commonly  to  North  Carolina, 
in  small  numbers  and  less  regularly  to  southern 
New  England,  and  perhaps  straying  around  the 
elbow  of  Cape  Cod  on  rare  occasions.  The  ten 
pounder  of  our  Atlantic  coast  is  represented  in 
tropical-warm  temperate  seas  in  various  other 
parts  of  the  world  by  relatives  so  close  that  they 
may  all  finally  prove  to  represent  only  the  one 
wide-ranging  species.13  Our  only  reason  for  men- 
tioning this  southern  fish  is  that  one  reported  as 
from  Chatham,  Mass.,  may  have  been  taken  on 
the  Gulf  of  Maine  shore  of  Cape  Cod.11  Ten 
pounders  are  taken  from  time  to  time  near  Woods 
Hole. 


'« Jordan  and  Evermann.  Bull.  47,  U.  8.  Nat.  Mus.,  Pt.  1,  1896,  p.  410. 
>'  Smith  (Sea  Fishes  Southern  Africa,  1949,  p.  86)  considers  this  probable. 
><  This  specimen,  taken  on  October  19, 1888,  and  reported  by  Bigelow  and 
8chroeder  (Copela,  1940,  p.  139)  Is  In  the  Museum  of  Comparative  Zoology. 


Figure  38. — Ten  pounder  (Elops  saurus),  Massachusetts.     From  Goode.     Drawing  by  H.  L.  Todd. 


FISHES   OF  THE  GULF   OF   MAINE 


87 


Tarpon  Tarpon  atlanticus  (Cuvier  and  Valenci- 
ennes) 1846 

Jordan  and  Evermann,  1896-1900,  p.  409,  fig.  177. 

Description. — The  tarpon  is  herring-like  in  gen- 
eral form  and  appearance,  but  it  is  made  easily 
recognizable  by  the  fact  that  the  last  ray  of  the 
dorsal  fin  is  greatly  prolonged,  its  free  portion 
being  as  long  as  the  fin  is  high  or  longer,  and  by 
the  presence  of  the  bony  plate  on  the  throat  men- 
tioned above  (p.  85)  in  the  characterization  of 
the  famdy  to  which  it  belongs.  Furthermore,  the 
anal  fin  of  the  tarpon  is  deeply  falcate;  that  of 
all  Gulf  of  Maine  herring-fishes  rhomboid  in  out- 
line. The  ventral  fins,  which  are  situated  under 
or  behind  the  dorsal  fin  in  herrings,  alewives,  shad, 
and  menhaden,  are  considerably  in  front  of  the 
dorsal  fin  in  the  tarpon,  while  the  lower  jaw  of  the 
latter  projects  relatively  further;  its  scales  are 
relatively  larger;  and  its  caudal  fin  is  relatively 
wider. 

Color.— Bright  silvery  all  over,  the  back  darker 
than  the  belly. 

Size. — Tarpon  grow  to  a  length  of  6  to  8  feet; 
the  longest  recorded  was  8  feet  2  inches;  the 


heaviest  taken  on  rod  and  reel  weighed  247 
pounds.15 

General  range. — Tropical  and  subtropical  coasts 
of  America,  from  Brazil  to  Long  Island,  casually  to 
Cape  Cod,  and  to  Nova  Scotia,  where  it  has  been 
recorded  off  Isaacs  Harbor  and  in  Harrigan 
Cove.18  Its  chief  center  of  abundance  is  in  the 
West  Indies,  about  Florida,  and  in  the  Gulf  of 
Mexico. 

Occurrence  in  the  Gulf  oj  Maine.- — A  specimen 
5%  feet  long,  taken  at  Provincetown  on  July  25, 
1915,17  is  the  only  record  of  the  tarpon  in  the  Gulf 
of  Maine,  which  it  reaches  only  as  an  accidental 
straggler  from  the  south. 

Round  herring  Etrumeus  sadina  (Mitchill)  1815 

Jordan  and  Evermann,  1896-1900,  p.  420. 

Description. — The  most  distinctive  feature  of 
this  fish,  among  herrings,  is  that  its  belly  is 
rounded,  not  sharp  edged.  It  is,  furthermore,  the 
most  slender  of  our  herrings,  its  body  being  only 

>•  Taken  on  rod  and  reel  in  the  Panuco  River,  Mexico,  Mar.  24, 1938,  by 
H.  W.  Sedgewick. 
i*  Halkett,  Check  List,  Fishes  Canada,  Newfoundland,  1913,  p.  46. 
i'  Radcliffe,  Copeia,  No.  26, 1916,  p.  3. 


Figure  39. — Tarpon  (.Tarpon  atlanticus),  New  Jersey.     From  Goode.     Drawing  by  H.  L.  Todd. 


Figure  40. — Round  herring  (Etrumeus  sadina). 


88 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


one-sixth  as  deep  as  long,  thus  suggesting  a  smelt 
in  its  general  outline.  Its  dorsal  fin,  too,  stands 
wholly  in  front  of  the  ventrals  instead  of  over  the 
latter,  as  in  herring,  alewives,  and  shad ;  and  there 
are  fewer  anal  fin  rays  (only  about  13,  whereas  the 
herring  has  about  17,  the  alewife  about  19,  and  the 
shad  about  21)  than  any  of  the  latter. 

Color. — Olive  green  above  with  silvery  sides  and 
belly. 

Size. — Eight  to  ten  inches  long  when  adult. 

General  range. — Atlantic  and  Gulf  of  Mexico 
coasts  of  the  United  States;  occasionally  common 
as  far  north  as  Woods  Hole;  sometimes  straying 
past  Cape  Cod,  to  the  mouth  of  the  Bay  of  Fundy. 

Occurrence  in  the  Gulf  of  Maine. — This  southern 
fish  has  been  taken  at  Provincetown,  Mass., 
whence  the  Museum  of  Comparative  Zoology  has 
two  specimens;  one  was  taken  in  the  Yarmouth 
River  which  empties  into  Casco  Bay,  and  one  in 
the  bay  itself  on  September  15,  1924;18  it  has  been 
reported  from  Jonesport,  Maine;  also  from  East- 
port,  Maine,  in  1908."  And  a  number  of  them 
Were  taken  at  Campobello  Island,  at  the  mouth  of 
Passamaquoddy  Bay  in  September  1937.20 

Herring  Clupea  harengus  Linnaeus  1758 

Sea    herring;    Labrador    herring;   Sardine; 
Sperling;  Brit 

Jordan  and  Evermann,  1896-1900,  p.  421. 

Description. — The  sea  herring  is  typical  of  its 
family  in  form,  with  body  so  flattened  that  it  is 
much  deeper  than  thick;  moderately  pointed  nose; 
large  mouth  situated  at  the  tip  of  the  snout  and 

'•  Reported  to  us  by  the  late  Walter  H.  Rich  of  the  U.  S.  Bureau  of  Fish- 
eries. 

"  Reported  in  the  newspapers. 

»  Reported  by  Leim,  Progr.  Rept.  21,  Atlantic  Biol.  Sta.  Fish.  Res.  Bd. 
Canada,  1937,  p.  5;  and  by  McKenzie,  Proe.  Nova  Scotia  Inst.  Sei.,  vol.  20, 
1939,  p.  15. 


lower  jaw  projecting  a  little  beyond  the  upper  when 
the  mouth  is  closed;  sharp-edged  belly;  and  deeply 
forked  tail.  The  dorsal  fin  stands  over  the  much 
smaller  ventrals,  its  origin  about  midway  the 
length  of  the  body.  The  scales  are  large,  their 
rear  margins  rounded,  and  so  loosely  attached  that 
they  slip  off  at  a  touch.  There  is  no  adipose  fin, 
and  its  absence  at  once  distinguishes  all  the  her- 
rings from  any  of  the  salmon  tribe.  The  chief  ana- 
tomical character  separating  the  sea  herring  from 
the  shad  and  from  the  several  alewives  (genus  Po- 
molobus)  is  that  it  has  an  oval  patch  of  small  teeth 
on  the  vomer  bone  in  the  center  of  the  roof  of  the 
mouth.  Conspicuous  field  marks  separating  her- 
ring from  shad,  hickory  shad,  and  alewife  are  that 
the  point  of  origin  of  its  dorsal  fin  is  about  midway 
of  the  length  of  its  trunk  (considerably  farther  for- 
ward in  the  others) ;  its  body  is  not  so  deep,  a  differ- 
ence shown  better  in  the  illustrations;  and  the 
sharp  midline  of  its  belly  is  only  very  weakly  saw- 
toothed  but  is  usually  strongly  so  in  the  others, 
especially  along  the  space  between  ventral  and 
anal  fins. 

Color. — Deep  steel  blue  or  greenish  blue  on  the 
back  with  green  reflections;  the  sides  and  belly 
silvery;  the  change  from  dark  belly  to  pale  sides 
often  marked  by  a  greenish  band.  The  gill  covers 
sometimes  glisten  with  a  golden  or  brassy  gloss; 
indeed,  fish  just  out  of  the  water  are  iridescent  all 
over  with  different  hues  of  blue,  green,  and  violet; 
but  these  colors  soon  fade,  leaving  only  the  dark 
back  and  silvery  sides.  The  ventral  and  anal  fins 
are  translucent  white;  the  pectorals,  however,  are 
dark  at  the  base  and  along  the  upper  edge;  the 
caudal  and  dorsal  fins  are  dark  grayish  or  shading 
into  green  or  blue. 

Size. — Herring  grow  to  a  length  of  about  17 
inches  and  to  a  weight  of  about  IK  pounds. 

Habits. — The  herring  is  a  fish  of  open  waters, 


Figure  41. — Herring  (Clupea  harengus).     From  Goode.     Drawing  by  H.  L.  Todd, 


FISHES   OF  THE  GULF  OF   MAINE 


89 


traveling  as  a  rule  in  schools  of  hundreds  or  thou- 
sands; single  fish  are  seldom  seen,  or  even  small 
companies  of  a  few  dozen.  As  a  rule  all  the  indi- 
vidual members  of  a  school  are  about  the  same 
size,  whether  large  or  small.  It  is  not  known  how 
long  any  given  school  may  preserve  its  identity  as 
such.  Fridriksson  and  Aasen,21  it  is  true,  found 
that  herring  tagged  and  released  together  might  be 
recaptured  from  widely  separated  localities,  sug- 
gesting that  schools  are  more  or  less  temporary 
formations.  But  this  may  not  apply  to  schools 
that  have  assembled  under  natural  conditions. 

When  a  school  is  at  the  surface,  as  often  hap- 
pens on  a  calm  day,  its  presence  is  betrayed  by  a 
fine  rippling  of  the  water,  but  we  have  never  seen 
herring  "finning"  or  lifting  their  noses  above  the 
surface  as  menhaden  often  do  (p.  114).  They 
come  to  the  surface  most  often  by  night,  when 
their  presence  is  betrayed  by  their  luminous  trails, 
if  the  water  is  "firing,"  as  we  have  often  seen.  A 
school  is  likely  to  be  more  or  less  stationary  when 
feeding,  its  members  swimming  slowly  to  and  fro 
and  drifting  as  a  whole  with  the  current.22  But 
at  other  times  schools  are  seen  traveling  with  in- 
dividual fish  swimming  side  by  side,  rank  below 
rank,  as  far  down  in  the  water  as  the  eye  can  see 
from  a  boat,  all  heading  in  one  direction  appar- 
ently with  some  purposeful  intent.  We  have 
often  watched  schools  of  "sardine"  size  streaming 
close  past  a  certain  rocky  headland  in  the  southern 
side  of  Massachusetts  Bay,  seemingly  in  unending 
procession. 

As  Dr.  Huntsman  points  out,  "There  is  no  in- 
dication that  herring  swim  against  the  current 
unless  the  water  is  somewhat  turbulent."  23  If 
they  do  so  under  such  conditions,  it  depends  on 
the  relationship  between  their  rate  of  swimming 
and  the  strength  of  the  current  whether  they  ac- 
tually make  headway  against  it  or  lose  ground, 
tail  first. 

We  might  also  add  that  schools  of  herring,  like 
schools  of  menhaden,  are  not  so  easily  frightened 
by  the  approach  of  a  boat,  as  mackerel  often  are, 
and  striped  bass.  Herring  do  not  jump  unless 
frightened.  But  the  smaller  sizes  are  often  seen 
jumping  when  pursued  from  below  by  larger  pred- 


sl  Rept.,  Norwegian  Fish.  Mar.  Invest.  SkriJter,  vol.  9,  No.  11,  Eept.  1, 
1950,  p.  22. 

»  Huntsman  (James  Johnstone  Memorial  Vol.,  1934,  p.  83)  gives  an  in- 
teresting account  of  the  movements  of  herring  schools  in  Passamaquoddy 
Bay. 

•»  James  Johnstone  Memorial  Vol.,  1934,  p.  84. 

210941—53 7 


atory  fishes,  such  as  silver  hake  or  striped  bass, 
a  common  spectacle.  Fridriksson  and  Aasen 
found  that  herring,  held  in  live-nets,  swam  con- 
stantly at  a  rate  of  about  0.2  to  0.25  sea  miles  per 
hour  (6-8  meters  per  minute)  when  not  disturbed. 
And  it  is  certain  that  they  are  capable  of  long 
journeys,  for  a  number  of  herring  tagged  on  the 
northeast  coast  of  Iceland  have  been  recaught  in 
southern  Norway,  and  some  vice-versa.24 

The  activity  of  the  herring  is  controlled  in  great 
part  by  the  temperature  of  the  water.  In  Pas- 
samaquoddy Bay,  for  example,  they  are  "ob- 
served to  move  very  sluggishly  when  the  water  is 
coldest  in  February  and  March,"  26  and  probably 
this  applies  all  around  the  periphery  of  our  Gulf, 
for  the  upper  20  fathoms  ordinarily  cools  to  about 
33  to  36°  F.  during  those  months,  with  the  sur- 
face often  chilling  to  the  freezing  point  of  salt 
water  in  bays  and  harbors.  The  herring  become 
active  again  when  the  water  has  warmed  to  about 
40  to  43°. 

Food. — The  herring  is  a  plankton  feeder.  When 
first  hatched,  and  before  the  disappearance  of  the 
yolk  sac,  the  larvae  (European)  feed  on  larval 
snails  and  crustaceans,  on  diatoms,  and  on 
peridinians,  but  they  soon  begin  taking  copepods, 
and  depend  exclusively  on  these  for  a  time  after 
they  get  to  be  12  mm.  long,  especially  on  the 
little  Pseudocalanus  elongatus.26  As  they  grow 
older  they  feed  more  and  more  on  the  larger 
copepods  and  amphipods,  pelagic  shrimps,  and 
decapod  crustacean  larvae.  Examination  of  1 ,500 
stomachs27  showed  that  adult  herring  near  East- 
port  were  living  solely  on  copepods  and  on  pelagic 
euphausiid  shrimps  (Meganyctiphanes  norwegica), 
fish  less  than  4  inches  long  depending  on  the  former 
alone,  while  the  larger  herring  were  eating  both. 

When  feeding  on  euphausiids,  we  have  often  seen 
them  pursuing  the  individual  shrimps,  which 
frequently  leap  clear  of  the  water  in  their  efforts 
to  escape.  Even  in  winter,  when  shrimp  are 
rarely  seen  at  the  surface,  Moore  found  them  an 
important  article  in  the  diet  of  the  Eastport 
herring.     And  it  is  likely  that  the  local  appear- 

«  Fridriksson  and  Aasen,  Rept.  Norwegian  Fish.  Mar.  Invest.,  Skrifter, 
vol.  9,  No.  11,  Rept.  1, 1950,  pp.  26-27. 

M  Huntsman,  James  Johnstone  Memorial  vol.,  1934,  p.  83. 

»  The  diet  of  herring,  young  and  old,  in  the  English  Channel  and  in  the 
North  Sea  has  been  described  by  Lebour  in  a  series  of  papers  (see  especially 
Jour.  Mar.  Biol.  Assoc.  United  Kingdom,  vol.  12, 1921,  pp.  458-467),  by  Hardy 
(British  Fisheries  Invest.,  Ser.  3,  vol.  7,  No.  3, 1924),  and  by  Jesperson  (Medd. 
Komm.  Havund.  Ser.  Plankton,  vol.  2,  No.  2,  1928,  Copenhagen). 

«  Moore,  Rept.  U.  S.  Comm.  Fish.  (1896),  1898,  p.  402. 


90 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


ances  and  disappearances  of  schools  of  large  fish  in 
the  open  Gulf  are  connected  with  the  presence  or 
absence  of  euphausiid  shrimp  of  one  species  or 
another.  A  few  of  the  larger  fish,  however,  as 
well  as  the  smaller  ones,  will  usually  be  found 
full  of  copepods,  even  when  both  shrimp  and 
copepods  abound,  and  copepods  are  the  chief 
dependence  of  all  our  herring,  large  and  small, 
in  the  absence  of  shrimp.  The  amphipod  genus 
Euthemisto  also  is  an  important  food  for  herring 
in  European  seas;  hence  the  absence  of  Euthemisto 
from  the  herring  stomachs  examined  by  Moore 
and  by  us  has  doubtless  been  due  to  the  com- 
parative scarcity  of  this  large  active  crustacean 
in  the  coastwise  waters  of  the  Gulf  of  Maine. 

The  particular  species  of  copepods  on  which 
Gulf  of  Maine  and  Woods  Hole  herring  depend 
have  not  been  identified,  but  we  might  guess  that 
Calanus  predominates,  with  Pseudocalanus,  Acar- 
tia,  and  Centropages,  and  Temora  also,  at  its  times 
of  abundance,  while  Euchaeta  offers  a  rich  food 
supply  when  the  schools  seek  the  deep  waters  of 
the  basin  frequented  by  these  mammoth  copepods. 

In  default  of  an  abundant  supply  of  Crustacea, 
and  sometimes  even  when  these  are  plentiful, 
herring  feed  on  whatever  molluscan  larvae,  fish 
eggs,  Sagittae,  pteropods,  annelids  that  the  water 
contains,  even  on  microscopic  objects  as  small  as 
tintinnids  and  Halosphaera.  But  the  smaller 
microscopic  plants,  either  diatom  or  peridinian, 
are  never  found  in  the  stomachs  of  herring  more 
than  15  to  20  mm.  long,  probably  because  their 
gill  rakers  are  not  fine  enough  to  retain  them. 

Although  herring  normally  are  not  fish  eaters, 
small  launce,  silversides,  and  the  young  of  their 
own  species  have  been  found  in  them  at  Woods 
Hole.  And  Templeman28  reports  them  as  con- 
suming quantities  of  small  capelin,  in  winter,  in 
Newfoundland  waters. 

Herring  ordinarily  pick  up  their  food  objects 
individually  by  a  "definite  act  of  capture"  as 
Battle  expresses  it,29  while  she  found  that  herring 
in  the  aquarium  at  St.  Andrews  did  not  feed  in 
complete  darkness,  though  they  did  in  faint  light. 
But  it  seems  that  when  feeding  on  very  small 
objects  they  may  strain  these  out  with  their 
branchial  sieves  as  the  manhaden  does  (p.  114),  for 
Moore,  a  very  accurate  observer,  described  them 
as   swimming   open   mouthed   when   feeding   on 

'■  Bull.  Newfoundland  Government  Lab.,  No.  17, 1948,  p.  133. 
»  Ann.  Kept.  Biol.  Board  Canada  (1933),  1934,  p.  14-15. 


minute  crustaceans,  crossing  and  recrossing  on 
their  tracks.30 

Doubtless  it  is  because  of  their  feeding  habits 
that  herring  seldom  take  a  baited  hook,  if  they 
ever  do.  But  we  think  it  likely  that  large  ones 
when  feeding  on  shrimp  would  take  an  artificial 
fly,  as  spent  and  hungry  alewives  will  (p.  104)  on 
their  return  to  salt  water,  and  as  shad  will  on  their 
way  upstream  (p.  109). 

Enemies. — The  herring  is  the  best  of  all  bait 
fishes  in  our  Gulf,  where  it  is  preyed  upon  by  all 
kinds  of  predaceous  fish,  especially  by  cod, 
pollock,  haddock,  silver  hake,  striped  bass,  mack- 
erel, tuna,  salmon,  and  dogfish,  and  by  the  mack- 
erel sharks.  Silver  hake,  in  particular,  often  drive 
schools  of  herring  up  on  our  beaches,  where 
pursued  and  pursuers  alike  strand  on  the  shoaling 
bottom.  We  once  saw  this  happen  at  Cohasset  in 
Massachusetts  Bay  many  years  ago,  on  an  October 
morning,  when  hake  and  herring  were  so  inter- 
mingled in  shallow  water  at  the  height  of  the  car- 
nage that  we  soon  filled  our  dory  with  the  two, 
with  our  bare  hands.  The  finback  whales  also 
devour  herring  in  great  quantities.  The  short- 
finned  squid  {Ilex)  likewise  destroys  multitudes 
of  the  young  sardines.  On  one  occasion  near 
Provincetown,  in  June  1925,  we  watched  packs  of 
perhaps  10  to  50  squids  circling  around  a  school 
of  2-  to  4-inch  herring,  bunching  them  into  a 
compact  mass.  Individual  squids  then  darted 
in,  seized  one  or  two  herring,  ate  only  a  small 
part,  then  darted  back  for  more.  A  silvery  streak 
of  fragments  of  dead  herring  remaining  along  the 
beach  bore  witness  to  the  carnage. 

Breeding  habits,  development  and  growth. — Much 
attention  has  been  devoted  to  the  breeding  habits 
and  growth  of  the  herring  by  European  zoologists, 
by  Moore,  and  by  Huntsman  in  our  own  Gulf, 
and  by  Lea  3X  in  more  northern  Canadian  waters. 

Herring  may  spawn  in  spring,  in  summer  or 
autumn,  according  to  locality,  or  both  in  spring 
and  autumn  (for  further  information  on  this 
matter,  see  p.  98).  They  do  so  chiefly  on  rocky, 
pebbly,  or  gravelly  bottoms,  on  clay  to  some  ex- 
tent, probably  never  on  soft  mud.  Spawning  in 
the  Gulf  of  Maine  (including  the  Bay  of  Fundy), 
takes  place  chiefly  from  2  or  3  fathoms  down  to 
about  30  fathoms;  perhaps  never  in  the  littoral 


W  Rept.  TJ.  S.  Comm.  Fish.  (1896),  1898,  p.  402. 
»  A  ge  and  growth  of  the  herrings  In  Canadian  waters. 
1914-16  (1919),  pp.  75-164. 


Oanad.  Fish.  Exped., 


FISHES   OF  THE  GULF  OF   MAINE 


91 


zone,  nor  has  herring  spawn  ever  been  reported  as 
cast  up  by  the  surf  on  the  beaches  of  New  England, 
a  fate  that  often  overtakes  it  in  the  Gulf  of  St. 
Lawrence.  Occasionally  they  spawn  as  deep  as 
100  fathoms  in  Scandanavian  waters,  perhaps  also 
in  the  eastern  basin  of  our  Gulf  where  the  sea  floor 
is  hard,  not  soft  and  muddy  as  it  is  in  the  basins  in 
the  western  side.  During  the  act  of  reproduction 
as  observed  by  Moore  at  Cross  Island  and  at 
Machias,  Maine,  "the  fish  were  darting  rapidly 
about,  and  those  who  have  opportunity  to  see  the 
fish  spawning  in  more  shallow  water  where  ob- 
servation is  more  favorable,  state  that  both  males 
and  females  are  in  constant  motion,  rubbing 
against  one  another  and  upon  the  bottom,  appar- 
ently by  pressure  aiding  in  the  discharge  of  the 
eggs  and  milt."  32 

A  female  herring  may  deposit  from  20,000  to 
upwards  of  40,000  eggs,  according  to  her  age  and 
size,  averaging  about  30,000.  In  sexually  mature 
herrings,  the  genital  organs  are  so  large  just  before 
spawning  commences  that  they  make  up  about 
one-fifth  the  total  weight  of  the  fish. 

The  eggs  sink  to  the  bottom,  where  they  stick 
in  layers  or  clumps  to  the  sand  or  clay,  to  sea- 
weeds, or  to  stones,  by  means  of  their  coating  of 
mucus,  or  to  any  other  objects  on  which  they 
chance  to  settle.  They  are  often  found  massed 
on  net  warps,  anchors,  and  anchor  ropes.  The 
individual  eggs  are  1  to  1.4  mm.  in  diameter,  de- 
pending on  the  size  of  the  parent  fish  and  also, 
perhaps,  on  the  local  race  of  fish  involved.  The 
period  of  incubation  is  governed  by  temperature; 
European  students  tell  us  that  it  requires  as  long 
as  40  days  at  38-39°,  15  days  at  44^6°  and  11 
days  at  50-51°  F.;  while  experiments  on  the  Mas- 
sachusetts coast  by  the  U.  S.  Bureau  of  Fisheries 
gave  10  to  12  days  in  the  temperature  prevailing 
there  in  autumn.  And  MacFarland  33  found  that 
all  of  the  eggs  kept  at  Grand  Manan  at  about  59  ° 
(15°  C.)  hatched,  but  that  none  hatched  at  32-41° 
(0-5°  C),  and  that  all  died  that  were  warmed  to 
68°  (20°  C.).  Ten  to  fifteen  days  might  be  stated 
as  an  average  incubation  period  for  the  Gulf  of 
Maine,   under  existing   temperatures. 

The  larvae  of  the  herring  family  are  very  slender 
and  can  easily  be  distinguished  from  all  other 
young  Gulf  of  Maine  fish  of  similar  form  (e.  g., 
launce,  smelt,  or  rock  eel)  by  the  location  of  the 

»  Moore,  Rept.  U.  S.  Comm.  Fish.  (1896),  1888,  p.  412. 
»  Rept.  Biol.  Board  Canada  (1930),  1931,  p.  23. 


Figure    42. — Eggs    of    the    herring    (Clupea    harengus), 
attached  to  seaweed  (European).     After  Ehrenbaum. 

vent,  which  is  so  far  back  that  it  lies  close  to  the 
base  of  the  tail.  But  it  requires  critical  examina- 
tion to  distinguish  our  several  clupeoids  one  from 
another  in  their  early  stages. 

The  sea  herring  is  about  5  to  6  mm.  long  at 
hatching,  with  a  small  yolk  sac  that  is  absorbed 
by  the  time  a  length  of  about  10  mm.  is  reached. 
The  dorsal  fin  is  formed  at  15  to  17  mm.;  the  anal 
at  about  30  mm.;  the  ventrals  are  visible  and  the 
tail  well  forked  at  30  to  35  mm. ;  and  at  about  40 
mm.  (1%  in.),  the  little  fish  begins  to  look  like  a 
herring. 

According  to  Huntsman's  observations,  fry 
produced  on  the  Grand  Manan  spawning  grounds 
in  late  summer  and  early  autumn  grow  to  a  length 
of  17  to  20  mm.  by  the  end  of  November  or  first 
of  December;  they  are  26  to  50  mm.  (1-2  in.)  long 
in  March  and  April  and  50  to  60  mm.  (2-2%  in.) 
long  by  June  when  fry  of  this  size  are  abundant 
in  the  St.  Andrews  region.  This  is  in  line  with 
our  own  observations  that  fry  of  2  to  2%  inches 
(50-65  mm.)  predominate  among  the  young  her- 
ring at  Provincetown  at  the  end  of  June,  and  fry 
of  2%  to  4  inches  (54  to  100  mm.)  on  Nantucket 


92 


FISHERY  BULLETIN   OF  THE    FISH   AND   "WILDLIFE   SERVICE 


Figure  43. — Larval  stages  of  the  herring  (Clupea  harengus) 
European.  After  Ehrenbaum.  A,  newly  hatched,  7 
mm.;  B,  10  mm.;  C,  19  mm.;  D.  29  mm.;  E,  young  fry, 
41  mm. 

Shoals  in  mid-July.  They  grow  to  about  3%  to 
near  5  inches  (90-125  mm.)  by  the  end  of  their 
first  year  of  life;  fish  of  that  size,  presumably  of 
the  previous  autumn's  hatch,  are  abundant  in  the 
fall  in  the  Bay  of  Fundy,  and  at  Boothbay,  Maine. 
The  growth  rate  is  about  the  same  at  Woods  Hole, 
where  herring  spawned  in  October  and  early  No- 
vember are  3  to  5  inches  (76-125  mm.)  long  by  the 
following  autumn.  The  Norwegian  herring,  also, 
average  about  5  inches  (125  mm.)  long  at  the  end 
of  their  first  year,  according  to  Hjort,  and  North 
Sea  herring  are  about  4  inches  (100  mm.)  long 
then.34 

Subsequent  growth. — The  herring  has  proved  a 
particularly  favorable  object  for  growth  studies 
based  on  the  structure  of  the  scales.36  Without 
pursuing  this  subject,  which  would  lead  us  far 
afield,  we  may  point  out  that  herring  not  only 
grow  at  different  rates  at  different  times  of  year, 
with  the  contrast  between  the  rapid  growth  of 
summer  and  the  slow  growth  of  the  winter  greater 
or  less  in  different  seas,  but  that  they  grow  rapidly 
when  young  and  slowly  thereafter  in  some  locali- 


ties, whereas  they  may  grow  slowly  at  first  in 
other  localities,  but  sustain  a  more  even  growth 
to  old  age. 

The  Dogger  Bank  herring,  for  example,  in  the 
North  Sea  approximate  4  inches  in  length  at  the 
end  of  the  first  year,  8%  to  9  inches  at  the  end  of 
the  third  year,  10%  at  the  end  of  the  sixth,  and  11% 
to  12  inches  at  the  end  of  the  ninth,  though  with 
considerable  variation.  The  Norwegian  herring, 
however,  spawned  in  the  year  1899,  averaged  only 
7%  inches  when  3  years  old,  but  were  as  large  as  the 
Dogger  Bank  fish  of  equal  ages  by  their  sixth  year 
and  subsequently.38  Newfoundland  herring  grow 
more  slowly  at  first  than  those  in  the  southern  side 
of  the  Gulf  of  St.  Lawrence,  but  catch  up  with 
them  as  they  grow  older. 

Huntsman  credits  the  Bay  of  Fundy  herring 
with  about  10  inches  at  the  end  of  their  third  year; 
i.  e.,  when  4  years  old,  which  agrees  closely  with 
an  average  growth  of  9%  inches  at  4  years  as  cal- 
culated by  Lea  for  Gulf  of  St.  Lawrence  fish.  The 
average  growth  rate  of  the  older  Bay  of  Fundy  fish 
probably  falls  between  that  of  the  Gulf  of  St. 
Lawrence  fish  and  that  of  the  herring  of  outer 
Nova  Scotia  which  grow  a  little  faster;  i.  e.,  to 
between  10%  and  11%  inches  at  5  years;  between 
11  and  12%  inches  at  7  years;  and  between  12%  and 
13%  inches  at  9  years.37  Bay  of  Fundy  herring 
make  most  of  their  growth  from  May  to  Septem- 
ber. In  the  southern  parts  of  our  Gulf,  where  the 
growth  period  probably  continues  a  month  later 
into  the  fall,  they  may  grow  as  fast  as  they  do  along 
outer  Nova  Scotia. 

When  the  little  herring  have  reached  an  age  of 
about  2  years  and  a  length  of  7%  to  8  inches 
(190-200  mm.)  they  accumulate  large  amounts  of 
fat  among  the  body  tissues  and  viscera  during  the 
warm  months  of  the  year  when  growing  rapidly, 
but  lose  this  fat  in  winter  and  also  at  the  approach 
of  sexual  maturity.  We  can  bear  witness  and 
the  fact  is  well  known  to  fishermen  that  this  "fat" 
stage  is  as  characteristic  of  American  waters  as  of 
European,  where  "fat"  herring  are  the  objects  of 
extensive  fisheries. 

According  to  Moore,  who  examined  thousands 


»  Huntsman  (Canad.  Fish.  Exped.  (1914-1915),  1919,  pp.  168-169)  believed 
he  could  recognize  spring  as  well  as  autumn-spawned  herring  fry  in  the  Bay 
of  Fundy,  and  credits  the  former  with  a  length  of  about  90  mm.  by  the  first, 
and  160  mm.  by  the  second,  winter.  But  this  seems  to  call  for  confirmation, 
it  being  unlikely  that  any  herring  now  spawn  there  in  spring  (p.  98). 

»  See  Lea  (Canad.  Fish.  Exped.  (1914-15),  1919,  pp.  75-164)  tor  an  account  of 
age  determination  by  analysis  of  the  scales,  as  applied  to  the  herring. 


'*  Rapp.  and  Proc.  Verb.,  Cons.  Intornat.  Erplor.  Mer,  vol.  20,  1944. 

»'  As  scaled  from  Lea's  diagrams  (Canad.  Fish.  Exped.  1914-1915  (1929), 
figs.  40  and  41).  It  has  been  found  that  the  Norwegian  herring  grow  from 
April  to  September  only,  remaining  practically  stationary  in  length  from 
October  until  March;  see  Lea  (Pub.  de  Circ,  Cons.  Perm.  Internat.  Explor. 
Mer,  No.  61,  1911,  pp.  35-57)  and  Hjort  fRapp.  Proc.  Verb.,  Cons.  Perm. 
Internat.  Explor.  Mer,  vol.  20, 1914). 


FISHES  OF  THE  GULF  OF   MAINE 


93 


of  fish  about  Eastport,  herring  rarely  spawn  when 
less  than  9%  inches  long;  usually  not  until  they 
are  10  to  10%  inches;  and  most  of  the  spawners 
are  12  to  13  inches  long.  This  means  that  some 
few  spawn  when  only  3  years  old,  if  the  growth 
schedule  outlined  above  is  correct,  but  most  of 
them  not  until  4  years  or  older,  to  continue 
spawning  annually  thereafter  as  long  as  they  live. 
In  Norwegian  waters,  too,  a  few  spawn  at  3  years, 
many  at  4  years,  and  the  majority  at  5  years;  some 
few  not  until  6  years  old.  Herring  have  been  seen 
as  old  as  20  years,  and  they  may  live  even  longer. 

Success  of  reproduction. — The  relative  abund- 
ance of  any  species  of  sea  fish  from  year  to  year 
depends  less  on  how  many  individuals  spawn  in 
any  locality  than  it  does  on  how  many  of  the 
resultant  fry  survive.  And  the  many  age  analyses 
that  have  been  made  of  herring  in  European  waters 
have  proved  that  while  a  very  large  crop  of  young 
may  be  produced  in  some  years,  hardly  any  are  in 
others,  even  in  favorable  nurseries.  Apparently 
this  applies  more  to  the  northern  breeding  grounds 
than  to  the  southern  (to  some  extent,  however,  to 
all)  the  result  being  that  the  herring  spawned  in 
some  one  favorable  breeding  season  may  dominate 
the  schools  over  large  areas  for  many  years,  or 
until  another  successful  breeding  year  comes, 
producing  another  large  crop.  In  Norwegian 
waters,  for  example,  the  herring  produced  in  1904 
was  dominant  in  the  catches  for  the  next  six  years, 
at  least;  this  is  a  classic  instance.  Lea  found, 
similarly,  that  herring  hatched  that  same  year 
(1904)  dominated  the  catches  on  the  west  coast  of 
Newfoundland  as  long  afterwards  as  1914  and 
1915.  And  while  precise  information  is  not  avail- 
able for  our  Gulf,  no  doubt  the  same  rule  governs 
there. 

One  case,  at  least,  is  well  documented  of  a 
particular  body  of  Bay  of  Fundy  herring  that 
received  no  important  recruitment  for  something 
like  10  years,  when  the  few  still  remaining  seem 
to  have  disappeared,  from  old  age  (p.  99). 

Various  explanations  have  been  proposed  to 
account  for  this,  such  as  abundance  or  scarcity  of 
microscopic  plankton,  favorable  or  unfavorable 
temperature,  salinity,  or  other  factors,  all  of 
which  may  enter  in.  And  while  it  is  during  the 
first  few  weeks  of  life  that  the  herring  is  most 
vulnerable,  it  is  also  possible  that  the  conditions 


under  which  the  parent  fish  lived  for  the  year 
preceding  spawning  may  influence  the  fate  of  the 
fry.  Whatever  the  explanation,  the  fact  that 
such  fluctuations  do  occur  from  year  to  year,  in 
the  numbers  of  fry  reared  is  of  the  greatest 
practical  interest  to  all  concerned  with  the  sea 
fisheries,  as  evidence  that  variations  existing  in 
the  stock  of  herring,  and  consequently  in  the 
catch,  may  be  due  more  to  the  success  or  failure 
of  reproduction  than  to  any  effect  the  fishery  may 
have  on  the  stock. 

General  range. — Both  sides  of  the  North  Atlan- 
tic. Off  the  European  coast  the  herring  ranges 
north  to  Norway,  Iceland,  Spitzbergen,  and  the 
White  Sea;  south  to  the  Straits  of  Gibraltar.  It 
is  known  on  the  American  coast  as  far  north  as 
northern  Labrador  and  the  west  coast  of  Green- 
land; regularly  and  commonly  as  far  south  as 
Cape  Cod  and  Block  Island ;  and  it  is  occasionally 
seen  in  small  numbers  as  far  south  as  Cape  Hat- 
teras  in  winter.  It  is  replaced  by  a  close  ally 
(C.  pallasii)  in  the  North  Pacific. 

Occurrence  in  the  Gulf  of  Maine. — To  fist  the 
localities  where  herring  have  been  recorded  would 
be  to  mention  every  hamlet  along  our  coasts 
whence  fishing  boats  put  out,  for  more  or  less 
herring,  large  or  small,  appear  at  one  season  or 
another  around  the  entire  coast  line  of  the  Gulf, 
and  on  the  offshore  fishing  banks  as  well.  They 
also  enter  bays  and  estuaries  freely,  but  they  have 
never  been  reported  in  our  Gulf  from  water  that 
is  appreciably  brackish;  perhaps  2.8  percent 
salinity  38  may  be  set  at  about  their  lower  limit. 

The  distribution  of  commercial  catches,  plotted 
by  Needier  (fig.  44)  39  shows  that  herring  are  far 
more  plentiful  from  Casco  Bay  eastward  along 
the  coast  of  Maine,  and  especially  in  the  Passama- 
quoddy  Bay-Grand  Manan  region  than  they  are 
along  the  western  shores  of  the  Gulf  on  the  one 
hand,  or  up  the  Bay  of  Fundy  on  the  other,  or 
along  western  Nova  Scotia.  Thus  the  landings 
per  unit  length  of  coast  averaged  3  times  as  great 
for  the  Passamaquoddy-Grand  Manan  region 
and  for  the  coast  of  Maine  to  Mount  Desert,  as 
for  the  coast  sector  from  Mount  Desert  past 
Penobscot  Bay;  about  4  times  as  great  as  for  the 
Maine  coast  as  a  whole,  westward  and  southward 

»  Surface,  in  Bay  of  Fundy  in  May. 

"  A  reliable  index,  for  the  herring  is  a  valuable  fish. 


94 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


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Figtjre  44. — Catches  of  herring  for  the  year  1919.     Each  dot  represents  400,000  pounds.     After  Needier. 


from  Penobscot  Bay;  and  13  times  as  great  as  for 
the  coast  of  Massachusetts,40  for  the  years  1919, 
1928,  1929,  and  1930.41 

Present  day  landings  of  upwards  of  30  million 
pounds  of  sardines  alone,  for  Charlotte  County, 
New  Brunswick,  even  in  poor  years,  up  to  some- 
thing like  100  million  pounds  in  good  years,  plus 
some  9-14  million  pounds  of  larger  herring,  con- 
trasted with  a  maximum  of  only  about  17  to  18 
million  pounds  reported  for  1947  for  any  sector  of 
the  Maine  coast  of  comparable  length,42  show  that 
the  Passamaquoddy-Grand  Manan  region  has  not 
lost  its  preeminence  as  a  herring  center.  The 
abundance  of  little  herring  there  is,  in  fact,  the 
outstanding  feature  of  the  distribution  of  fishes  in 

*•  Omitting  the  landings  for  Suffolk  County,  Mass.,  since  these  represent 
flsh  discharged  at  Boston  by  the  vessel  fisheries  from  offshore. 

«  Graham,  Jour.  Biol.  Board  Canada,  vol.  2,  No.  2,  1936,  p.  129,  table  2. 

«  Scattergood  has  given  an  interesting  analysis,  regional  and  seasonal,  of 
the  1947  catch  of  herring  for  the  coast  of  Maine. 


the  Gulf  of  Maine.  A  catch  of  about  2,400,000 
pounds  for  Massachusetts  in  1947,  contrasted 
with  some  11,300,000  pounds  for  the  Penobscot 
Bay  region  alone  in  that  year,  illustrates  how  much 
less  rich  in  herring  the  southwestern  side  coast  line 
of  the  Gulf  is  than  the  sector  that  happens  to  be 
the  least  productive  part  of  the  northern  coast 
line  of  the  Gulf. 

Fishermen  tell  us,  too,  that  herring  are  much 
more  regular  in  their  occurrence  from  year  to  year 
in  the  Passamaquoddy-Grand  Manan  region  than 
they  are  either  off  western  Nova  Scotia  in  the  one 
direction,  or  along  the  coast  of  Maine  in  the  other. 
And  this  is  borne  out  by  such  statistics  as  are  con- 
veniently available.  Thus  only  one-fourth  to  one- 
fifth  as  many  pounds  of  herring  were  caught  in  the 
Penobscot  Bay  region  43  in  1947  as  either  eastward 


»  Scattergood's  statistical  areas  11-14. 


FISHES   OF  THE  GULF  OF   MAINE 


95 


to  Lubec  Narrows  on  the  one  hand,  or  westward 
past  Casco  Bay  to  Cape  Elizabeth  on  the  other,** 
whereas  the  catches  for  1919  were  rather  evenly 
distributed  along  the  northern  and  eastern  Maine 
coast  as  a  whole. 

We  find  herring  even  more  and  more  sporadic  in 
their  appearances  and  disappearances,  both  from 
place  to  place,  from  week  to  week,  and  from  year  to 
year,  passing  southward  around  the  western 
periphery  of  the  Gulf.  Very  few,  for  example,  are 
seen  on  the  southern  side  of  Massachusetts  Bay  in 
some  years  (as  in  1950  and  1951);  many  schools  in 
others.  And  herring  are  such  wandering  fish  in 
general,  here  today  and  gone  tomorrow  even  in 
their  centers  of  abundance,  that  the  successful 
location  of  the  weirs  depends  largely  on  intimate 
local  knowledge  and  on  close  observation  of  the 
movements  of  the  schools. 

Herring  appear,  also,  to  be  far  less  plentiful  on 
the  offshore  banks  and  less  regular  in  their  occur- 
rences there  than  they  are  in  their  inshore  center 
of  abundance  in  the  northeastern  part  of  the  Gulf. 
Trawlers,  it  is  true,  occasionally  pick  up  schools  on 
Georges  Bank  and  on  Browns  Bank,  as  in  1931, 
when  catches  of  3,000  pounds  were  reported  on  the 
northern  edge  of  Georges  and  of  2,800  pounds  on 
the  southwestern  part  in  October.  Schools,  too, 
are  occasionally  reported  as  seen  at  the  surface,  by 
Albatross  III  for  instance,  in  April-May  1950. 
Fishermen  used  sometimes  to  set  drift-nets  on 
Georges  for  herring  for  bait  in  the  days  of  the 
long  line  fishery,  and  small  numbers  up  to  130-160 
per  haul,  were  trawled  by  Albatross  III,  widespread 
on  the  western  part  in  depths  of  20  to  50 
fathoms  in  May  1950,  as  well  as  off  southern  New 
England.46  But  it  is  more  usual  for  trawlers 
operating  on  Georges  to  pick  up  only  odd  fish  or 
none.  Thus  the  maximum  catch  on  any  trip 
during  the  otter  trawl  investigation  of  1913  was 
only  a  dozen  or  two;  42  hauls  by  the  Eugene  H, 
in  late  June  1951,  yielded  only  one  herring,  fishing 
from  Nantucket  Lightship  out  onto  the  south- 
central  part  of  Georges;  and  the  stomachs  of  cod 
caught  on  Georges  seldom  contain  herring,  if 
they  ever  do.46 

The  appearance  of  schools  of  large  herring  or  of 
small  is  distinctly  a  seasonal  event  off  most  parts 

44  Coast  sectors  of  comparable  length. 

«  Average  catches  per  haul  about  56  fish  at  22  to  40  fathoms,  and  28  at  41  to 
50  fathoms,  but  only  6  at  51  to  60  fathoms. 

«  W.  F.  Clapp  found  no  herring  in  many  cod  and  haddock  stomachs 
examined  by  him  on  Qeorges  Bank. 


of  our  coast,  and  the  picture  is  made  still  more 
complex  by  differences  in  the  behavior  of  sardine- 
size,  "fat,"  and  spawning  herring,  the  reasons  for 
which  are  not  yet  well  understood. 

The  newly  spawned  fry,  less  than  %  of  an  inch 
(9-11  mm.)  long,  have  been  taken  in  September 
in  the  lower  part  of  the  Bay  of  Fundy,  a  product, 
doubtless,  of  the  Grand  Manan  and  West  Nova 
Scotia  spawning;  also  in  October  in  Gloucester 
Harbor  where  one  tow-net  haul  yielded  us  a  great 
number  on  the  24th,  in  1916.  And  they  are  to  be 
expected  wherever  herring  spawn  in  numbers  in 
any  particular  year.  It  seems  likely  that  most  of 
them  remain  near  their  birth  place  during  their 
first  autumn  and  winter,  when  the  circulation  of 
the  Gulf  is  in  its  least  active  stage.  But  they 
become  widely  distributed  during  the  spring 
(March-May),  when  1%  to  2  inches  (30-50  mm.) 
long,  both  in  the  lower  Bay  of  Fundy,  around  the 
entire  periphery  of  the  open  Gulf,  east  as  well  as 
west,  out  over  the  basin,  and  on  the  northern  and 
eastern  parts  of  Georges  Bank.47 

Little  seems  to  be  known  in  detail  about  the 
movements  of  herring  during  their  first  year,  but 
those  that  find  their  way  into  enclosed  waters 
where  mid-summer  temperatures  are  high,  such  as 
Duxbury  and  Plymouth  Bays  and  Provincetown 
Harbor,  appear  to  move  out  during  the  early  part 
of  the  summer,  being  reported  as  far  less  plentiful 
there  in  June  than  they  are  in  April  and  May. 
Sardine-size  herring,  4  to  8  inches  long  including 
1-  and  2-year-olds,  are  to  be  expected  in  abundance 
all  summer  east  of  Penobscot  Bay,  and  particularly 
in  the  Passamaquoddy  Bay  region,  where  they 
support  the  sardine  fishery  for  which  the  latter 
is  famous,  and  where  they  are  present  throughout 
the  year. 

It  is  probable,  however,  though  not  proved, 
that  the  1-  to  2-year-olds  (fish  in  their  second  and 
third  years)  do  not  appear  along  the  southwestern 
coasts  of  the  Gulf  until  several  months  later  in 
the  season  than  the  little  fish  of  %  to  2  inches  do, 
that  were  hatched  the  preceding  autumn.  Thus 
it  usually  is  not  until  late  June,  July,  or  August 
that  "sperling"  of  4  to  7  inches  are  reported  in 
numbers  off  the  Massachusetts  coast,  or  that  we 

«  During  March  and  April  1920  we  took  them  near  Cashes  Ledge,  on  the 
northern  and  eastern  parts  of  Qeorges  Bank,  off  Seal  Island;  off  Yarmouth, 
Nova  Scotia;  near  Macblas,  Maine,  and  over  the  basin  in  the  offing;  near 
Boothbay;  and  near  the  Isles  of  Shoals.  Graham  (Jour.  Biol.  Board  Canada, 
vol.  2,  No.  2,  1936,  p.  112,  fig.  8)  found  them  equally  widespread  in  the  open 
Gulf  in  May  1932,  also  in  the  lower  Bay  of  Fundy  (none,  however,  at  the  head 
of  the  Bay). 


96 


FISHERY  BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


have  seen  them  there.  They  are  even  more  er- 
ratic, too,  in  their  appearances  and  disappearances 
in  Massachusetts  Bay  and  along  Cape  Cod  than 
they  are  to  the  eastward  of  Mount  Desert.  At 
Cohasset,  for  example,  on  the  southern  shore  of 
the  Bay  where  we  have  had  many  years'  experi- 
ence, schools  of  sperling  are  here  today  in  summer 
and  early  autumn,  but  gone  tomorrow.  It  is 
also  our  impression  that  the  sperling,  like  the 
larger  herring,  are  not  only  far  less  concentrated 
in  favorable  localities  around  the  southwestern 
shore  of  the  Gulf  than  they  are  to  the  north  and 
east,  but  far  less  numerous  on  the  whole.*8 

These  first  two  year  classes  (the  fish  in  their 
first  year  having  grown  to  a  length  of  3  or  4  inches 
by  September;  those  in  their  second  year  to  7  to 
9  inches)  begin  to  thin  out  from  the  shore  waters 
of  the  open  Gulf  after  the  middle  of  October  as 
the  water  cools,  and  few  "sardines"  are  taken 
there  after  early  December. 

The  corresponding  ebb  and  flow,  so  to  speak, 
for  the  sardine  is  suggested  in  a  striking  way  by 
the  average  monthly  catches  of  sardines  by  the 
weirs  in  Charlotte  Co.,  New  Brunswick  (Passama- 
quoddy  Bay,  Campobello,  and  Grand  Manan) 
for  the  year  1920,  which  are  equally  illustrative 
of  conditions  today : 


Month 

January 

February.. 
March 


Founds  Month  Founds 

11,000  July 3,315,000 

None  August 6,  475,  000 

56,000  September 6,730,000 

April 1,049,000  October 6,012,000 

May 3,036,000  November 1,325,000 

June. 2,542,000  December 147,000 

Here,  however,  the  seasonal  variation  (as  Dr. 
Huntsman  informs  us)  is  simply  a  matter  of  local 
availability,  for  sardines  remain  in  Passama- 
quoddy  Bay  all  winter,  but  do  not  move  about 
much  then.  Probably  the  sardines  winter  mostly 
on  the  bottom.  And  there  is  no  reason  to  sup- 
pose that  the  bulk  of  them  travel  far  in  any  part 
of  the  Gulf. 

Very  little  is  known  about  the  Gulf  of  Maine 
herring  during  their  third  summer,  when  they 
have  passed  the  "sardine"  or  sperling  stage  and 
have  not  yet  reached  spawning  age.  In  some 
years  these  "fat"  herring,  as  they  are  often  called, 


or  "summer"  herring,  weighing  up  to  about  one 
pound  (they  are  called  "spawn"  herring  locally, 
but  this  is  an  error),  are  taken  in  the  traps  at 
Provincetown  for  a  week  or  so  about  mid-April; 
they  are  taken  at  about  the  same  time  off  Glouces- 
ter (in  1915  they  were  reported  8  to  15  miles 
off  Cape  Ann  on  the  17th),  and  they  are  said  by 
the  fishermen  to  "show"  first  off  Seguin  Island  in 
May  and  June,  off  Mount  Desert  late  in  summer. 
Doubtless  they  form  a  large  part  (just  what  pro- 
portion is  not  known)  of  the  catches  of  herring 
larger  than  sardines  that  are  made  in  the  Passa- 
maquoddy  Bay  region,  also  around  Grand  Manan. 
As  a  rule  few  of  them  are  taken  inside  the  inner 
islands  elsewhere,  though  they  came  into  the  har- 
bor of  Boothbay  about  May  14  in  1914. 

When  a  mackerel  seiner  picks  up  a  school  of 
herring  out  in  the  open  Gulf  in  summer,49  or  when 
an  otter  trawler  makes  a  catch  of  herring  on 
Georges  Bank  (p.  95),  most  of  them  are  very  fat 
and  show  no  signs  of  approaching  sexual  matu- 
rity. Thus  it  seems  that  they  tend  to  keep  farther 
offshore  than  do  either  the  younger  herring  or  the 
still  older  mature  herring. 

The  peak  season  for  herring  larger  than  "sar- 
dines" inshore  in  the  northeastern  part  of  the 
Gulf  is  ordinarily  from  July  through  October;  i.  e., 
some  2  months  less  than  that  for  the  sardines  (see 
p.  96).  But  a  greater  proportion  of  the  larger 
fish  continue  available  there  through  the  cold 
months  than  of  the  younger  fish,  to  judge  from 
the  fact  that  considerably  larger  catches  are  made 
of  big  herring  in  winter  than  of  sardines,  whereas 
the  total  local  catch  is  much  larger  for  the  latter 
than  for  the  former. 

A  report  w  on  the  average  monthly  landings  of 
large  herring  for  Charlotte  County,  for  the  period 
1920-1931,  to  the  nearest  1,000  pounds,  follows: 


Month 

January 

February. . 
March 


48  No  particular  attention  is  paid  to  sperling  around  Massachusetts  Bay, 
for  they  are  too  small  to  be  In  demand  for  bait,  and  they  are  not  plentiful 
enough  (or  not  concentrated  enough)  to  support  a  sardine  fishery  there. 


Pounds  Month  Pounds 

132,000  July. 1,065,000 

164,000  August 4,334,000 

275,000  September 7,098,000 

April....  312,000  October 2,817,000 

May 306,  000  November 646,  000 

June 284,000  December 268,000 

Large  herrings,  yearly  average 17,  701,  000 

"Sardines",  yearly  average 30,  698,  000 

«  Many  events  of  this  sort  have  been  reported.  For  example,  a  large  catch 
of  fat  summer  herring  was  made  on  Georges  Bank  and  reported  to  the  Massa- 
chusetts Commissioners  in  the  mid-summer  of  1901. 

»  From  Graham,  Jour.  Biol.  Board  Canada,  vol.  2,  No.  2,  1936,  p.  130, 
table  3. 


FISHES  OF  THE  GULF  OF   MAINE 


97 


Large  herring  (Dr.  Hunstman  tells  us)  are  also 
present  there  throughout  late  winter  and  spring, 
though  few  find  their  way  then  into  the  weirs. 

In  the  southeastern  part  of  the  Gulf,  as  typified 
by  Cape  Cod  Bay,  large  herring  appear  inshore  in 
greatest  numbers  to  about  June  and  again  in  the 
autumn,  with  very  few  (and  not  many  sardines) 
in  June  or  July.  This  is  illustrated  by  the  largest 
and  smallest  catches  made  in  8  traps  at  North 
Truro  for  different  months  during  the  years  1946 
to  1952.  The  following  data  are  contributed  by 
the  Pond  Village  Cold  Storage  Company: 

Minimum  Maximum 

Month  (in  pounds)  (in  pounds) 

April 0  117,375 

May 221  623,550 

June - -. -.  0  88,657 

July 0  0 

August 0  1,000 

September.. 0  57,287 

October 0  9,526 

November 0  176,435 

The  earliest  catch  of  sardines  there  in  those  years, 
or  in  1935,  1938,  or  1943  was  sometime  in  May, 
the  latest  November  16  to  17;  the  earliest  catcb 
of  large  herring  was  made  between  April  20  and 
30,  the  latest  on  December  10th. 

In  most  years  the  large  herring  vanish  from  the 
Massachusetts  coast  at  some  time  in  December. 
In  1950,  for  example,  they  vanished  about  De- 
cember 4th  from  Ipswich  Bay,  where  considerable 
catches  had  been  made  for  some  time  previous  by 
about  15  boats.81 

Nothing  is  known,  definitely,  as  to  their  sea- 
sonal appearances  and  disappearances  over  the 
offshore  banks. 

About  all  that  is  known  of  the  movements  of 
the  large  mature  herring  (in  their  fourth  summer 
and  older)  is  that  they  are  encountered  in  num- 
bers only  for  the  brief  period  before,  during,  and 
after  the  spawning  season,  when  they  are  seen 
schooling  at  the  surface,  and  are  caught  along 
shore.  Fishermen  report  that  they  show  about 
the  off-lying  islands  some  time  before  they  make 
their  way  up  the  bays ;  two  or  three  weeks  earlier, 
for  instance,  at  Grand  Manan,  Jonesport,  and 
about  Mount  Desert  Island  than  within  Machias 
Bay.  They  are  said  to  appear  some  time  after 
the  middle  of  July  at  Isle  au  Haut  at  the  eastern 
entrance  of  Penobscot  Bay,  and  at  Castine  within 
the  Bay,  though  not  until  the  end  of  that  month 


11  This  happening  was  reported  in  the  daily  papers. 
210941—53 8 


or  the  first  of  August  at  Matinicus  Island.  Such 
of  them  as  visit  the  Massachusetts  Bay  region  are 
not  expected  there  until  the  last  week  in  Septem- 
ber. But  they  are  in  full  force  on  all  the  spawning 
areas  along  the  shores  of  the  Gulf  by  October, 
from  Grand  Manan  to  Cape  Cod;  they  are  equally 
widespread,  if  less  abundant,  inshore  in  November, 
and  they  are  reported  in  December  occasionally, 
and  even  later.  It  is  probable  that  as  the  fish 
spawn  out  most  of  them  move  out  promptly  from 
the  spawning  grounds  into  deeper  water,  for  fish 
recently  spent  are  not  often  reported  as  taken  in 
the  weirs. 

Probably  the  spawners  merely  descend  into 
deeper  water  to  winter,  as  is  the  case  in  European 
waters.  How  deep  the  great  body  of  them  go  is 
not  known.  But  is  has  been  proved  that  herring 
of  all  ages  remain  in  the  open  Bay  of  Fundy 
throughout  the  cold  season;  also  in  the  passages 
between  the  inner  and  outer  divisions  of  Passama- 
quoddy  Bay,  even  when  water  temperatures  there 
are  as  low  as  32°  F.62  And  the  abundance  of 
pelagic  euphausiid  shrimps  (a  favorite  herring 
food)  in  the  deeper  water  layers  of  the  northeast 
corner  of  the  Gulf  suggests  this  as  a  rich  winter 
pasture  for  them. 

Studies  carried  out  from  the  Atlantic  Biological 
Station  at  St.  Andrews,  chiefly  under  Dr.  A.  G. 
Huntsman's  M  leadership,  and  by  the  International 
Passamaquoddy  Fisheries  Commission64  during 
the  early  1930's  seem  to  us  to  have  proved  that 
the  factor  chiefly  responsible  for  the  great  concen- 
tration of  young  herring  in  the  Passamaquoddy 
region,  and  for  their  availability  to  the  weir  fishery 
there,  is  the  differential  circulation  of  the  shoaler 
and  deeper  water  layers  that  is  set  in  motion  by 
the  inflow  of  fresh  water  from  the  tributary 
streams  combined  with  superficial  currents  set  up 
temporarily  by  the  wind.  In  other  words,  the 
sardine-sized  herring  acts  as  does  any  planktonic 
animal  such  as  the  euphausiid  shrimps  and  the 
copepod  crustaceans,  on  which  it  feeds,  as  it 
swims  to  and  fro,  i.  e.,  it  drifts  with  the  current. 
In  technical  language,  it  is  "denatant." 

The  case  is  not  so  clear  for  the  larger  herring, 
not  because  there  is  any  reason  to  suppose  they 
can  direct  their  journeys  more  intelligently,  and 
because  any  directive  swimming  they  may  carry 

u  Huntsman.  James  Johnstone  Memorial  Vol.,  1934,  p.  82. 
•»  For  summary,  see  Huntsman,  James  Johnstone  Memorial  Vol.,  1934, 
pp.  95-96. 
«  See  Graham,  Jour.  Biol.  Board  Canada,  vol.  2, 1936,  No.  2,  pp.  93-140. 


98 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE    SERVICE 


out  is  far  more  effective  because  more  rapid;  but 
because  so  little  is  known  as  to  journeys  any  indi- 
vidual school  actually  makes  as  season  follows 
season,  whether  of  fat  herring  or  of  spawners. 
Perhaps  the  most  interesting  question  of  all,  and 
one  as  yet  unsolved,  is  how  and  why  the  spawning 
fish  seek  their  spawning  grounds  year  after  year, 
when  their  sex  organs  mature. 

Spawning  grounds  and  season. — It  appears  that 
the  most  productive  spawning  ground  for  our 
Gulf  formerly  was  and  still  is  at  the  mouth  of  the 
Bay  of  Fundy,  particularly  on  the  shoals  south- 
west of  Grand  Manan.  The  Trinity  Ledges  off 
western  Nova  Scotia  are  another  important 
ground;  and  herring  are  reported  as  spawning 
commonly,  though  irregularly,  in  Machias  Bay; 
about  Jonesport;  at  Mount  Desert;  in  French- 
mans  Bay;  among  the  islands  at  the  mouth  of 
Penobscot  Bay  (Swans,  Isle  au  Haut,  and  Matin- 
icus);  in  Casco  Bay;  also  about  Wood  Island  a 
few  miles  south  of  Cape  Elizabeth,  which  has  long 
been  known  as  the  resort  of  breeding  schools. 
Herring  have  also  been  found  spawning  off  the 
beaches  along  the  western  shore  of  the  Gulf, 
Ipswich  Bay,  for  example;  about  Cape  Ann;  in 
Massachusetts  Bay;  about  Provincetown;  along 
outer  Cape  Cod ;  in  the  Woods  Hole  region ;  near  No 
Mans  Land;  and  about  Block  Island  which  is  the 
southern  breeding  limit.  But  whatever  spawning 
does  take  place  either  southward  from  the  vicinity 
of  Cape  Elizabeth  on  the  one  hand,  or  in  the  inner 
parts  of  the  Bay  of  Fundy  on  the  other,  is  trifling 
as  compared  with  the  production  along  the 
eastern  coast  of  Maine  and  in  the  Grand  Manan 
region. 

Spawning  takes  place  both  along  shore  in  our 
Gulf  and  on  the  various  shoals  and  ledges  that  lie 
for  5  to  25  miles  off  the  coast,  a  habit  betrayed  by 
the  eggs  that  are  found  sticking  to  the  anchor 
ropes  of  fishing  vessels.  But  we  find  no  definite 
record  of  herring  spawning  on  Browns  or  Georges 
Banks,  nor  are  young  fry  known  there,  a  fact  that 
was  commented  upon  by  Storer  long  ago. 

Spawning  season.  —  Both  spring  -  spawning 
schools  and  summer-fall  spawning  schools  of 
herring  were  reported  formerly,  in  the  Bay  of 
Fundy,  the  spring  spawners  visiting  the  south 
(Nova  Scotia)  side  of  the  bay  from  Bier  Island 
at  the  mouth  in  as  far  as  Digby  Gut,  also  the 
Parrsboro  region  on  the  New  Brunswick  shore 
near  the  head  of  the  bay,  spawning  during  April 


and  May.  But  they  seem  never  to  have  been 
very  numerous,  and  it  is  not  known  whether  any 
spawn  now  in  the  bay  before  summer.  Spring- 
spawning  as  well  as  autumn-spawning  herring 
have  also  been  reported  to  us  by  fishermen  along 
the  west  coast  of  Nova  Scotia,  though  we  have 
not  been  able  to  verify  this.  Other  than  this, 
spring  spawners  are  neither  recorded  nor  rumored 
anywhere  in  the  Gulf  of  Maine. 

Around  Grand  Manan  and  in  Machias  Bay 
nearby,  the  heaviest  runs  of  summer-autumn 
spawners  usually  come  in  July,  August,  and  Sep- 
tember, the  spawning  season  continuing  until  late 
in  the  fall  in  some  years,65  but  not  commencing 
until  early  August,  and  ending  by  early  Oc- 
tober in  others.66  Passing  westward  we  find  the 
breeding  period  progressively  later  and  shorter; 
mid-August  for  example  until  October  around 
Petit  Manan  and  near  Mount  Desert,  while 
the  few  herring  that  spawn  farther  south  do  so 
chiefly  during  October  in  Ipswich67  and  Massa- 
chusetts Bays;  in  late  October  and  early  Novem- 
ber in  the  vicinity  of  Woods  Hole. 

So  many  observations  have  been  taken  in  the 
Gulf  from  the  vessels  of  the  Bureau  of  Fisheries, 
and  in  the  Bay  of  Fundy  by  the  Biological  Board 
of  Canada,  that  it  is  possible  to  establish  the 
temperatures  rather  closely  at  which  herring 
spawn  in  our  waters.  Around  Grand  Manan  and 
in  the  northern  part  of  the  Gulf  generally,  prac- 
tically all  spawning  is  carried  out  in  water  of  about 
46-52°  F.  But  such  herring  as  spawn  in  the 
southern  part  of  Massachusetts  Bay  and  along 
the  shores  of  Cape  Cod,  where  autumnal  cooling 
of  the  surface  waters  is  not  so  rapid  as  it  is  farther 
north,  may  do  so  in  slightly  warmer  water,  say 
up  to  53°  or  55°.  The  Gulf  of  Maine  herring 
spawn  in  rather  low  salinities  (such  characterize 
the  coastal  zone  as  a  whole  as  compared  with  the 
North  and  Norwegian  Seas),  the  most  saline  water 
in  which  it  is  known  to  spawn  within  our  limits 
being  not  saltier  than  33  per  mille,  the  freshest 
probably  about  31.9  per  mille.  They  never  spawn 
in  brackish  water  within  the  limits  of  the  Gulf, 
although  known  to  do  so  at  the  mouths  of  certain 
European  rivers  in  water  that  is  nearly  fresh. 

Destruction  by  natural  causes. — The  herring  is  a 
very  "tender"  fish,  prone  to  wholesale  destruction 

"  So  described  by  Moore,  Rept.  U.  S.  Comm.  Fish.  (1896),  1898,  p.  408. 

69  Dr.  Huntsman  informs  us  that  this  was  the  case  in  1917. 

«'  Allen,  Mem.  Boston  Soc.  Nat.  Hist.,  vol.  8,  No.  2, 1916,  p.  201. 


FISHES   OF  THE  GULF  OF   MAINE 


99 


both  by  stranding  on  beaches  during  storms,  and 
by  pollution  of  the  water.     Many  instances  of 
this  kind  have  been  reported.     Allen,68  for  exam- 
ple, saw  young  herring  in  windrows  for  miles  on 
the  strand  at  Rye  Beach,  N.  H.,  in  August  1911. 
A  slaughter  of  herring  (still  more  instructive  be- 
cause the  exact  course  of  events  was  followed)  took 
place  at  Cohasset,  on  the  south  shore  of  Massa- 
chusetts Bay,  in  October  1920.     On  the  5th  of 
that  month  a  large  school  of  "sperling,"  4  to  5 
inches  in  length,  ran  up   the  harbor   (which  is 
nearly  landlocked),  probably  driven  in  by  silver 
hake    (at   least   so   local   fishermen   said);   were 
trapped  there  by  the  falling  tide,  and  stranded  on 
the  mud.     So  numerous  were  they  that  the  flats 
were  entirely  covered  with  them  and  it  was  esti- 
mated that  20,000  barrels  of  fish  perished.     Dur- 
ing the  next  few  days  the  fish  (alternately  covered 
and  uncovered  by  the  tide)  decayed,  and  despite 
the  tidal  circulation,   so  fouled  the  water  that 
lobsters  impounded  in  floating  cars  died.     On  the 
10th  there  was  a  second  smaller  run  of  herring, 
and  on  the  15th  a  third  run  came  as  numerous  as 
the  first,  the  newcomers  dying  soon  after  they 
entered  the  harbor.     Altogether,  it  was  estimated 
that  50,000  barrels  of  fish  perished,  of  which  more 
than  90  percent  were  "sperling,"  5  to  10  percent 
were  large  adults,  and  a  few  were  small  mackerel 
and  silver  hake,  besides  large  numbers  of  smelt. 
The  flats  were  silvery  with  herring  scales  at  low 
tide  by  the  last  half  of  October,  when  we  saw 
them,  and  the  residents  about  the  harbor  found  the 
stench  almost  unbearable.     But  the  fish  decom- 
posed and  the  water  purified  itself  during  the 
winter  months. 

Mass  destructions  of  young  herring  have  also 
been  reported  in  other  Gulf  of  Maine  harbors. 
Thus,  Dr.  Austin  H.  Clark  reported  that  early  in 
August  1925  the  mud  flats  in  Manchester  Harbor, 
on  the  north  side  of  Massachusetts  Bay,  were 
white  with  stranded  herring  3  to  5  inches  long, 
packed  several  deep  at  low  tide  along  the  sides  of 
the  little  drains  and  hollows.  Another  such 
destruction  took  place  in  the  same  harbor  in  the 
summer  of  1928.  Vast  quantities  of  herring  spawn 
are  likewise  cast  up  on  the  beaches  every  year  to 
perish  in  north  European  waters;  this  also  happens 
to  some  extent  in  the  Gulf  of  St.  Lawrence. 

Numerical  abundance  and  importance. — Moore 


■  Mem.  Boston  Soo.  Nat.  Hist.,  vol.  8,  No.  2, 1916,  p.  202. 


(1898),  who  sifted  many  sources  of  information 
concluded  (we  believe  rightly)  that  no  general 
decrease  had  taken  place  in  the  abundance  of 
young  herring  at  the  mouth  of  the  Bay  of  Fundy 
up  to  that  time.  But  it  is  common  knowledge 
among  fishermen  that  both  the  numbers  visiting 
any  given  locality  on  our  coast  and  the  duration 
of  their  stay  varies  widely,  not  only  from  year  to 
year,  but  over  longer  periods.  Local  spawning 
grounds,  too,  may  be  abandoned  for  a  term  of 
years — a  common  occurrence.69 

The  best  documented  case  of  local  disappear- 
ances from  a  previously  productive  ground  took 
place,  as  Dr.  Huntsman  writes  us,60  from  the 
shoals  southwest  of  Grand  Manan,  whence  large 
herring  (previously  very  plentiful)  withdrew  in 
1877,  to  reappear  in  1881  on  the  Nova  Scotia 
coast  between  Cape  Sable  and  Digby.  Dr. 
Huntsman  has  suggested  that  they  had  circled 
the  Gulf  offshore,  for  their  exodus  from  the  Grand 
Manan  shoals  was  not  accompanied  by  any  coin- 
cident increase  in  the  catch  along  the  eastern  part 
of  the  coast  of  Maine,  but  rather  by  the  reverse.61 
They  persisted  on  the  Nova  Scotia  shore  until 
1890,  when  they  gave  out,  probably  from  old  age, 
for  the  large  herring  that  remained  in  the  Quoddy 
region  also  dwindled  in  numbers  as  shown  by  the 
collapse  of  the  winter  fishery  there,  evidence  that 
this  particular  body  of  herring  did  not  receive  any 
significant  recruitment  after  about  1880-1881. 
It  remains  to  be  seen  whether  large  herring  will 
ever  reappear  in  their  former  plenty  on  the  Grand 
Manan  ground,  as  they  did  about  1857  in  Massa- 
chusetts Bay,  where  the  stock  had  been  at  a  low 
ebb  since  1837;  or  whether  the  yearly  drain  on 
the  population  of  young  herring  by  the  sardine 
fishery  (well  started  by  about  1880)  is  too  great. 

The  largest  reported  catch  of  herring  for  the 
Gulf  as  a  whole  for  any  year  since  1928  for  which 
statistics  are  readily  available  was  219,131,500 
pounds  taken  in  1946,  divided  as  follows:  Massa- 
chusetts, 2,049,000  pounds;  Maine,  80,107,400 
pounds;  and  the  Canadian  shores  of  the  Gulf, 
136,975,100  pounds.  The  smallest  catch  was 
70,519,886  pounds  in  1932,  divided  5,687,254 
pounds,  3 1 ,988, 132  pounds,  and  32,844,500  pounds, 

»  Moore,  Rept.  U.  S.  Comm.  Fish.  (1896),  1898,  p.  430. 

•°  Based  on  Canadian  fishery  statistics. 

•i  Earll  (Fisheries  and  Fish.  Ind.  U.  S.,  sect.  5,  vol.  1,  1887,  pp.  423,  424) 
states  that  the  fishery  declined  near  Bois  Bubert  Island  from  1875  to  1880, 
and  that  the  catch  was  "considerably  below  average"  at  Matlnicus  during 
the  10  years  previous  to  1879. 


100 


FISHERY  BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


respectively.  In  1947,  incidentally,  the  herring 
catch  of  the  Gulf  was  topped  only  by  the  catches 
of  haddock  and  of  rosefish. 

It  is  not  clear  to  what  extent  this  range  in  the 
catch  from  year  to  year  is  due  to  fluctuations  in 
the  supply  offish;  to  differences  in  their  availability; 
or  to  the  sundry  economic  factors  that  enter  in. 
What  is  certain  is  that  with  some  80  percent  of 
the  catch  consisting  of  sardine-sized  fish  weighing 
only  about  one-half  ounce,  the  toll  taken  cannot 
have  been  less  than  1%  billion  fish  in  the  poorest 
of  recent  years,  5  to  6  billion  in  the  year  when  the 
yield  was  greatest,  i.  e.,  numbers  far  greater  than 
that  for  any  other  Gulf  of  Maine  fish.  Come  good 
year  then  or  bad,  Capt.  John  Smith's  account  of 
the  herring  of  our  Gulf  thus  applies  equally  well 
today:  "The  savages  compare  the  store  in  the  sea 
with  the  hair  of  their  heads,  and  surely  there  are 
an  incredible  abundance  upon  this  coast."  62 

The  sardine  catch  of  the  Bay  of  Fundy  is  made 
almost  wholly  in  weirs,  347  of  which  were  operated 
on  the  New  Brunswick  shore  in  1947.  On  the 
Maine  coast,  as  a  whole,  a  little  less  than  two- 
thirds  the  catch  of  herring,  large  and  small,  is 
made  in  weirs  and  in  purse  seines,  combined,  a 
little  more  than  one-third  nowadays  in  stop  seines 
(about  44,500,000  pounds  in  1947).  These  are 
used  "to  prevent  the  exit  of  the  herring  school  from 
a  cove  or  inlet.  .  .  .  The  seine  is  stretched  around 
the  school  with  the  ends  of  the  net  made  fast  to  the 
shore."  And  stop  seines  are  used  mostly  at  night, 
when  the  presence  of  fish  is  betrayed  by  their 
luminous  trails,  if  the  water  is  firing,  or  by  the 

•>  General  Hlstorle  of  Virginia,  New  England,  and  the  Summer  Isles,  1616, 
reprinted  In  1819  from  London  edition  of  1629,  p.  188. 


noise  they  make  as  they  "flip"  at  the  surface.*3 
And  some  are  still  caught  in  floating  traps  (about 
2  million  pounds  in  1947)  which  we  have  often 
seen  used  in  the  harbors  of  Mount  Desert. 

In  1947,  seemingly  a  representative  year,  purse 
seines  yielded  the  Maine  fishermen  nearly  as  much 
herring  (about  36,100,000  pounds)  as  the  weirs. 
Their  presence  is  detected,  Scattergood  tells  us, 
either  by  the  firing  of  the  water  if  by  night,  by  echo 
souDding  apparatus,  or  by  the  use  of  a  thin  wire 
suspended  in  the  water,  the  vibrations  of  which 
indicate  the  presence  of  fish  that  strike  it.  In  1947 
eleven  purse  seiners  were  active  in  the  fall  fishery 
for  Maine  herring.  How  many  were  engaged  in 
the  New  Brunswick  and  Maine  winter  fishery  is 
not  known. 

Large  catches  of  herring  when  on  bottom  also 
are  made  by  special  otter  trawls  in  European  wa- 
ters; and  of  the  closely  allied  herring  of  British 
Columbia  of  late.  But  the  possibility  of  develop- 
ing an  otter-trawl  fishery  for  herring  in  the  Gulf  of 
Maine  has  not  yet  been  explored. 

Finally,  we  may  remark  that  herring  fresh  from 
the  water  are  among  the  most  delicious  of  our 
fishes,  especially  the  small  sizes.  Their  only  draw- 
back is  that  they  do  not  keep  well,  being  rich- 
meated  and  oily,  and  in  the  larger  sized  fish  the 
many  hair-like  bones  are  troublesome. 

Hickory  shad  Pomolobus  mediocris  (Mitchill)  1815 
Fall  herring;  Shad  herring 

Jordan  and  Evermann,  1896-1900,  p.  425. 


•»  See  Scattergood,  U.  S.  Fish  and  Wildlife  Service,  Sp.  Scl.  Rept.  No.  67, 
1949,  p.  8,  for  further  details. 


Figtjbb  45. — Hickory  shad  (Pomolobus  mediocris) ,  Chesapeake  Bay  region  specimen. 

Todd. 


From  Goode.     Drawing  by  H.  L. 


FISHES   OF  THE  GULF   OF   MAINE 


101 


Description.— The  hickory  shad  differs  rather 
noticeably  from  the  sea  herring  in  that  the  point 
of  origin  of  its  dorsal  fin  is  considerably  in  front  of 
the  mid-length  of  its  trunk;  in  its  deep  belly  (a 
hickory  shad  13}£  in.  long  is  about  4  in.  deep  but  a 
herring  of  that  length  is  only  3  in.  deep) ;  in  the  fact 
that  its  outline  tapers  toward  both  snout  and  tail 
in  side  view  (fig.  15);  and  in  that  its  lower  jaw 
projects  farther  beyond  the  upper  when  its  mouth 
is  closed;  also,  by  the  saw-toothed  edge  of  its  belly. 
Also,  it  lacks  the  cluster  of  teeth  on  the  roof  of  the 
mouth  that  is  characteristic  of  the  herring.  One 
is  more  likely  to  confuse  a  hickory  shad  with  a  shad 
or  with  the  alewives,  which  it  resembles  in  the 
position  of  its  dorsal  fin,  in  the  great  depth  of  its 
body,  in  its  saw-toothed  belly  and  in  the  lack  of 
teeth  on  the  roof  of  the  mouth.  But  it  is  marked 
off  from  all  of  these  by  its  projecting  lower  jaw. 
There  is  also  a  small  difference  in  outline,  its  head 
tapering  more  to  the  snout,  as  seen  in  side  view 
(fig.  45) .  It  has  only  about  half  as  many  gill  rakers 
(19  to  21  on  the  lower  limb  of  the  first  gill  arch) 
as  either  the  alewife  or  the  blueback ;  and  its  upper 
jaw,  reaching  back  only  about  as  far  as  opposite 
the  center  of  its  eye,  is  shorter  than  that  of  the 
shad  in  which  it  reaches  as  far  as  the  rear  edge  of 
the  eye. 

Under  favorable  circumstances  its  color,  also, 
is  characteristic,  for  it  is  faintly  marked  on  the 
sides  with  dusky  longitudinal  stripes,  and  the  tip 
of  its  snout  is  dusky. 

Size. — This  is  the  largest  of  our  anadromous 
herrings  next  to  the  shad,  growing  to  a  length  of 
2  feet.  A  fish  about  15  inches  long  weighs  a 
pound,  one  of  18  inches,  2  pounds. 

Habits. — Nothing  is  known  of  the  habits  of  the 
hickory  shad  in  the  sea  to  differentiate  it  from  its 
close  relatives  of  the  herring  tribe  except  that  it  is 
more  of  a  fish  eater.  Launce,  anchovies,  cunners, 
herring,  scup,  silversides,  and  other  small  fish, 
squid,  fish  eggs,  and  even  small  crabs  have  been 
found  in  the  stomachs  of  hickory  shad  at  Woods 
Hole,  as  well  as  sundry  pelagic  Crustacea.  It 
will  strike  a  small  spinner  or  other  artifical  lure, 
and  it  gives  a  good  fight  when  hooked.  In  the 
southern  parts  of  its  range  it  is  described  as  running 
up  fresh  streams,  with  the  alewives  in  late  winter 
and  early  spring  to  spawn.94  But  it  appears  not 
to  do  so  in  the  streams  tributary  to  Chesapeake 

M  Smith  (N.  C.  Geol.  Econ.  Surv;  vol.  2,  1897,  p.  121)  describes  It  as  doing 
so  In  the  streams  tributary  to  Pamlico  Sound,  N.  C,  where  It  is  plentiful. 


Bay,  though  it  is  found  in  practically  all  of  them. 
This  opens  the  interesting  possibility  that  the 
"green"  fish  found  in  Chesapeake  Bay,  leave  the 
Bay,  perhaps  to  spawn  in  salt  water.66 

General  range. — Atlantic  coast  of  North  America 
from  the  Bay  of  Fundy  to  Florida. 

Occurrence  in  the  Gulf  of  Maine.- — The  hickory 
shad  is  a  southern  fish,  with  the  Gulf  of  Maine  as 
the  extreme  northern  limit  to  its  range.  It  is 
recorded  in  scientific  literature  only  at  North 
Truro;  at  Provincetown;  at  Brewster;  in  Boston 
Harbor;  off  Portland;  in  Casco  Bay;  and  from  the 
mouth  of  the  Bay  of  Fundy  (Huntsman  doubts 
this  record) ,  and  it  usually  is  so  uncommon  within 
our  limits  that  we  have  seen  none  in  the  Gulf 
ourselves.  But  in  1932  anglers,  trolling  for  striped 
bass  and  mackerel  off  the  Merrimac  River,  met  a 
run  of  hickory  shad.66 

It  is  much  more  plentiful  west  of  Cape  Cod, 
being  common  from  spring  throughout  summer  and 
early  autumn  at  Woods  Hole,  where  as  many  as 
3,500  have  been  taken  at  a  single  lift  of  one  trap. 
In  1919  the  Massachusetts  catch  of  hickory  shad, 
practically  all  from  the  south  coast,  amounted  to 
12,800  pounds,  and  none  are  listed  for  Massa- 
chusetts for  any  subsequent  year. 

Alewife  Pomolobus  pseudoharengus  (Wilson)  1811 
[approximate  date] 

Gaspereau;  Sawbelly;  Kyak;  Branch  herring; 
Fresh- water  herring;  Grayback 

Jordan  and  Evermann,  1896-1900,  p.  426. 

Description. — The  alewife  is  distinguishable  at 
a  glance  from  the  sea  herring  by  the  greater  depth 
of  its  body,  which  is  three  and  one-third  times  as 
long  as  deep  (an  alewife  of  13K  inches  is  about 
4  inches  deep;  a  herring  that  long  has  a  depth  of 
only  3  inches)  also  by  the  position  of  its  dorsal 
fin,  the  point  of  origin  of  which  is  considerably 
nearer  to  the  tip  of  the  snout  than  to  the  point  of 
origin  of  the  central  rays  of  the  tail  fin.  Further- 
more, the  alewife  is  much  more  heavily  built 
forward  than  the  herring,  and  the  serrations  on 
the  midline  of  its  belly  are  much  stronger  and 
sharper  (hence  the  local  name  "sawbelly"),  so 
much   so   that   a   practiced   hand   can   separate 

-  Hildebrand  and  Schrocder,  Bull.,  U.  S.  Bur.  Fish.,  vol.  43,  1928,  p.  84. 

M  The  Museum  of  Comparative  Zoology  received  one  from  this  run  from 
Dr.  J.  O.  Phillips,  caught  by  him  off  the  northern  end  of  Plum  Island,  October 
2,  1932. 


102 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  46. — Alewife  (Pomolobus  pseudoharengus) ,  Chesapeake  Bay  region  specimen. 

Todd. 


From  Goode.     Drawing  by  H.  L. 


herring  from  alewives  in  the  dark.  The  most 
useful  distinctions  between  the  alewife  and  the 
blueback  are  that  in  the  former  the  eye  is  broader 
than  the  distance  from  its  forward  edge  to  the  tip 
of  its  snout  and  the  back  grayish  green,  while  in 
the  latter  the  eye  is  only  about  as  wide  as  the 
distance  from  front  of  eye  to  tip  of  snout,  and  the 
back  is  dark  blue  (p.  107).  Also  the  lining  of  the 
abdominal  cavity  is  pale  grayish  or  pinkish  white 
in  the  alewife,  but  is  usually  dusky  or  blackish  in 
the  blueback.  But  this  distinction  may  not  hold 
in  all  cases. 

Alewives  are  distinguishable  from  young  shad 
by  their  smaller  mouths  with  shorter  upper  jaws; 
also  by  the  fact  that  the  lower  jaw  of  the  alewife 
projects  slightly  beyond  the  upper  when  the 
mouth  is  closed,  and  by  the  outline  of  the  edge  of 
the  lower  jaw,  the  forward  part  of  which  is  deeply 
concave  in  the  alewife  but  nearly  straight  in  the 
shad.  The  lack  of  teeth  on  the  roof  of  the  mouth 
distinguishes  the  alewife,  with  its  brethren  the 
hickory  shad  (p.  100)  and  blueback  (p.  106)  from 
the  sea  herring,  anatomically. 

Color. — The  alewife,  like'  the  herring,  is  grayish 
green  above,  darkest  on  the  back,  paler  and 
silvery  on  sides  and  belly.  Usually  there  is  a 
dusky  spot  on  either  side  just  behind  the  margin 
of  the  gill  cover  (lacking  in  the  herring)  and  the 
upper  side  may  be  faintly  striped  with  dark  longi- 
tudinal lines  in  large  fish.  The  sides  are  iridescent 
in  life,  with  shades  of  green  and  violet.  The  colors 
change,  to  some  extent,  in  shade  from  darker  to 
paler,  or  vice  versa,  to  match  the  bottom  below, 
as  the  fish  run  up  stream  in  shallow  water. 


Size. — The  alewife  grows  to  a  length  of  about  15 
inches,  but  adults  average  only  about  10  to  11 
inches  long  and  about  8  to  9  ounces  in  weight; 
16,400,000  fish  taken  in  New  England  in  1898 
weighed  about  8,800,000  pounds. 

Habits. — The  alewife,  like  the  shad  and  the 
salmon  makes  its  growth  in  the  sea,  but  enters 
fresh  water  streams  to  spawn.  This  "anadro- 
mous"  habit,  as  it  is  called,  forced  itself  on  the 
attention  of  the  early  settlers  on  our  coasts.  In 
the  words  of  an  eyewitness,  "experience  hath 
taught  them  at  New  Plymouth  that  in  April  there 
is  a  fish  much  like  a  herring  that  comes  up  into 
the  small  brooks  to  spawn,  and  when  the  water 
is  not  knee  deep  they  will  presse  up  through  your 
hands,  yea,  thow  you  beat  at  them  with  cudgels, 
and  in  such  abundance  as  is  incredible."  67  And 
they  are  no  less  persevering  in  their  struggles 
upstream  today.  Numbers  of  them  are  to  be 
seen  in  many  streams,  any  spring,  alternately 
swimming  ahead ;  resting  in  the  eddy  behind  some 
irregularity  of  the  bottom;  then  moving  ahead 
again,  between  one's  feet  if  one  happens  to  be 
standing  in  midstream.  And  they  are  much  more 
successful  than  the  shad  in  surmounting  fishways 
of  suitable  design.  During  the  early  runs  some- 
times one  sex  predominates,  sometimes  the  other, 
but  the  late  runs  consist  chiefly  of  males,  as  a 
rule,  and  these  are  said  to  outnumber  the  females 
greatly  on  the  spawning  grounds.  We  have  no 
firsthand  observations  to  contribute  on  this  score. 

Alewives  are  decidedly  general  in  their  choice 

■  Capt.  Charles  Whitbome,  In  "The  True  Travels  of  Oapt.  John  Smith," 
1S16,  vol.  2,  p.  260. 


FISHES   OF  THE   GULF   OF   MAINE 


103 


of  streams,  running  indifferently  up  rivers  as  large 
as  the  St.  John,  Merrimac  and  Potomac,  or 
streams  so  small  that  one  can  almost  leap  across, 
and  only  a  few  inches  deep.  In  large  rivers  they 
run  far  upstream — how  far  they  may  do  so  we 
do  not  know — or  their  journey  may  be  one  of 
only  a  few  yards,  as  it  is  in  the  artificial  cuts  that 
are  kept  open  through  barrier  beaches  to  allow 
the  fish  access  to  fresh  water  ponds  behind  the 
latter. 

The  alewife  spawns  in  ponds,  including  those 
back  of  barrier  beaches  (if  there  are  openings  to 
the  sea,  natural  or  artificial)  and  in  sluggish 
stretches  of  streams,  never  in  swift  water,  each 
female  depositing  from  60,000  to  100,000  eggs  or 
more,  according  to  her  size.68  Spawning  lasts 
only  a  few  days  for  each  group  of  fish. 

The  spent  fish  run  down  stream  again  so  soon 
after  spawning  that  many  of  them  pass  others 
coming  up,  as  we  have  often  seen;  fish  on  their 
return  journey  to  salt  water  are  familiar  sights  in 
every  alewife  stream. 

The  adults,  when  entering  streams  to  spawn, 
make  the  change  from  salt  water  to  fresh  within 
a  short  time  without  damage;  this  is  equally  true 
of  the  spent  fish  on  their  return  to  the  estuaries. 
But  Dr.  Huntsman  informs  us  that  they  appear 
unable  to  endure  repeated  changes  between  salt 
water  and  fresh,  and  that  great  numbers  are 
killed  in  this  way  in  the  estuaries  under  certain 
conditions  of  tide.  The  strain  of  spawning  leaves 
them  very  thin,  but  they  recover  rapidly  after 
they  reach  salt  water.  We  have  seen  spent 
alewives  that  had  already  put  on  considerable 
fat,  taken  from  a  trap  at  Provincetown  as  early 
in  the  season  as  July  16  (in  1915). 

Spawning  ordinarily  takes  place  at  tempera- 
tures of  about  55  to  60°.  The  eggs  are  about 
0.05  inches  in  diameter,  pink  like  those  of  the  sea 
herriDg,  and  they  stick  to  brush,  stones,  or  any- 
thing else  they  may  settle  upon.89  Incubation 
occupies  about  6  days  at  60°.  The  young 
alewives,  which  are  about  5  mm.  long  when 
hatched,  growing  to  15  mm.  when  a  month  old, 
soon  begin  to  work  their  way  downstream.  They 
have  been  seen  descending  as  early  as  June  15  in 
the  more  southerly  of  Gulf  of  Maine  streams; 

M  The  average  number  of  eggs  In  644  females  taken  In  the  Potomac  was 
102.800  (Smith,  N.  C.  Qeol.  and  Econ.  Survey,  vol.  2,  1907.    p.  123). 

"  The  development  of  the  eggs,  larval  stages,  and  young  fry  are  described 
by  Ryder  (Report,  U.  S.  Comm.  of  Fish.  (1885),  1887,  p.  505)  and  by 
Prince  Contr.  Canad.  Biol.  (1902-1905),  1907,  p.  96). 


successive  companies  of  fry  move  out  of  the  pond 
and  down  with  the  current  throughout  the 
summer;  and  by  autumn  the  young  alewives  have 
all  found  their  way  down  to  salt  water  when  2  to 
4  inches  long.  We  have  seined  young  alewives 
as  long  as  4  to  4K  inches  (102-115  mm.)  in  salt 
water  near  Seguin  Island,  Maine,  at  the  end  of 
July,  but  others,  only  3  to  3%  inches  long  (78-92 
mm.),  near  Mt.  Desert  Island  as  late  as  the  first 
of  October.  Thenceforth  the  alewife  lives  in 
salt  water  until  sexual  maturity. 

Hildebrand  and  Schroeder70  found  that  little 
alewives  in  Chesapeake  Bay  had  grown  to  about 
i)i  to  5  inches  long  by  the  time  they  were  1  year 
old. 

The  rate  of  growth  of  the  older  alewives,  in  salt 
water,  has  not  been  traced.  But  experiments  in 
planting  adult  alewives  in  ponds  in  which  there 
were  none  before,  led,  long  ago,  to  the  conclusion 
that  they  became  sexually  mature  at  3  or  4  years 
of  age,  for  none  of  their  progeny  returned  until 
3  or  4  years  after  the  original  plant.  Specific 
instances,  cited  by  Belding  71  are: 

(1)  Three  years  after  a  large  number  of  alewives 
were  hatched  in  Keene's  Pond,  Maine,  tributary 
to  the  Calais  River,  from  a  "plant"  of  mature  fish, 
a  run  of  adult  fish  entered  Keene's  Pond  stream 
where  none  had  ever  been  seen  before;  this  case  was 
reported  by  the  U.  S.  Bureau  of  Fisheries.  (2) 
The  establishment  of  a  fishery,  in  the  same  way  at 
Plymouth,  Mass.,  in  4  years  after  restocking  in 
1865;  and  (3)  G.  M.  Besse  obtained  results  in 
3  years  in  ponds  in  Wareham,  Mass. 

The  fact  that  alewives  have  been  known  to 
return,  for  spawning,  to  streams  in  which  their 
parents  had  been  planted,  lends  support  to  the 
"parent  stream"  theory;  i.e.,  that  alewives,  like 
shad,  tend  to  spawn  in  the  stream  system  in 
which  they  were  hatched.  But  a  much  more 
intensive  study  is  needed  of  this  interesting 
question  before  any  categorical  statement  can  be 
made,  as  to  how  generally  this  is  true;  and  to 
what  extent  their  return  depends  on  their  never 
having  wandered  far  afield. 

Food. — The  alewife  is  chiefly  a  plankton  feeder 
like  the  herring;  copepods,  amphipods,  shrimps, 
and  appendicularians  were  the  chief  diet  of  speci- 
mens examined  by  Vinal  Edwards  and  by  Linton 

'•  Bull.  U.  S.  Bur.  Fish.,  vol.  43, 1928,  p.  91i 

"  Rept.  Alewife  Fish.  Mass.,  Mass.  Dept.  Conservation,  Div.  Fish,  and 
Game,  1921,  p.  18. 


104 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


at  Woods  Hole.  However,  they  also  take  small 
fish,  such  as  herring,  eels,  laun.ce,  cunners,  and 
their  own  species,  as  well  as  fish  eggs.  Unlike 
herring,  alewives  often  contain  diatoms  even  when 
adult.  Alewives  fast  when  they  are  running  up- 
stream to  spawn,  but  when  the  spent  fish  reach 
brackish  water  on  their  return  they  feed  ravenously 
on  the  shrimp  that  abound  in  the  tidal  estuaries  and 
which  they  can  be  seen  pursuing.  We  have  often 
hooked  alewives  on  an  artificial  fly  at  such  times. 

Movements  at  sea. — The  alewife  is  as  gregarious 
as  the  herring,  fish  of  a  size  congregating  in  schools 
of  thousands  of  individuals  (we  find  record  of 
40,000  fish  caught  in  one  seine  haul  in  Boston 
Harbor)  and  apparently  a  given  school  holds 
together  during  most  of  its  sojourn  in  salt  water. 
But  they  are  sometimes  caught  mixed  with  men- 
haden, or  with  herring.  Alewives,  immature  and 
adult,  are  often  picked  up  in  abundance  in  weirs 
here  and  there  along  the  coast,72  and  it  is  likely 
that  the  majority  remain  in  the  general  vicnity  of 
the  fresh  water  influence  of  the  stream-mouths 
and  estuaries  from  which  they  have  emerged,  to 
judge  from  the  success  of  attempts  to  strengthen 
or  restore  the  runs  of  alewives  in  various  streams, 
mentioned  above.  But  it  is  certain  that  some  of 
them  wander  far  afield,  for  catches  of  up  to  3,000 
to  4,000  pounds  per  haul  were  made  by  otter 
trawlers  some  80  miles  offshore,  off  Emerald 
Bank,  Nova  Scotia  (lat.  about  43°  15'  N.,  long, 
about  63°  W.)  at  60  to  80  fathoms,  in  March 
1936.73 

Odd  alewives  were  reported  from  Georges  Bank 
and  the  South  Channel  in  March,  June,  August, 
and  November  of  1913.  Some  (up  to  78  per 
haul)  were  trawled  by  Albatross  III  about  25  to  60 
miles  out  off  southern  New  England  in  May  1950; 
also  18  adults,  10  to  11  inches  long,  70  odd  miles 
off  Barnegat,  N.  J.,  on  March  5,  1931 ;  and  we  saw 
60  alewives  trawled  at  the  25-fathom  line  off 
Marthas  Vineyard 7i  in  late  June,  1951  by  the 
Eugene  H.  Where  these  wanderers  come  to  shore 
to  spawn,  if  they  succeed  in  doing  so  at  all,  is  an 
interesting  question. 

It  seems  likely  from  various  lines  of  evidence 
that  alewives  tend  to  keep  near  the  surface  for 
their  first  year  or  so  in  salt  water,  and  while  they 


«  Huntsman  (Contr.  Canad.  Biol.,  [1921]  1922,  p.  58)  reports  its  young  at 
Campobello  Island,  Bay  of  Fundy,  in  December  and  March. 

*•  Reported  by  Vladykov,  Copeia,  1936,  No.  3,  p.  168.  One  vessel  brought 
in  about  10,000  pounds. 

'<  At  lat.  40°  68'  N.;  long.  70°  32*  W. 


are  inshore  when  older.  But  practically  nothing 
is  known  as  to  the  depths  to  which  they  may 
descend  if  (or  when)  they  move  offshore,  there 
being  no  assurance  that  those  taken  by  trawlers 
were  not  picked  up,  while  the  trawls  were  being 
lowered  or  hauled  up  again. 

General  range. — Gulf  of  St.  Lawrence  and  north- 
ern Nova  Scotia  south  to  North  Carolina,  running 
up  into  fresh  water  to  spawn;  landlocked  races 
also  exist  in  Lake  Ontario,  in  the  Finger  Lakes  of 
New  York,  and  in  certain  other  fresh-water  lakes.7' 

Occurrence  in  the  Gulj  of  Maine. — When  the 
white  man  crossed  the  Atlantic  probably  there 
was  no  stream  from  Cape  Sable  to  Cape  Cod  but 
saw  its  annual  run  of  alewives  unless  they  were 
barred  by  impassable  falls  near  the  mouth. 

And  while  its  numbers  have  declined  during  the 
past  two  centuries  and  its  range  has  been  restricted, 
both  by  actual  extirpation  from  certain  streams  by 
overfishing,  by  the  pollution  of  the  river  waters  by 
manufacturing  wastes,  and  by  the  erection  of  dams 
that  it  cannot  pass,  the  alewife  is  a  familiar  fish 
still,  all  along  around  our  coast 78  and  yields  an 
abundant  catch  in  many  of  our  streams.  Ale- 
wives are  taken  commonly  about  Yarmouth,  Nova 
Scotia;  in  the  Annapolis  Basin;  in  Minas  Channel; 
and  farther  still,  up  the  Bay.  Alewives  still  run 
in  most  of  the  streams  tributary  to  the  Bay  of 
Fundy,  many  in  the  St.  John.  A  few  are  taken  in 
the  weirs  in  Passamaquoddy  Bay ;  while  young  ones 
have  been  taken  around  Campobello  Island;  as 
deep  as  50  fathoms.  They  enter  the  large  river 
systems  all  along  the  coasts  of  Maine  and  New 
Hampshire,  likewise  many  small  streams,  the  re- 
quirements being  that  these  shall  lead  to  ponds  or 
have  deadwaters  of  sufficient  extent  along  their 
courses,  and  no  dams  or  falls  that  the  alewives 
can  not  surmount.  At  Boothbay  Harbor,  for  in- 
stance, a  considerable  number  of  alewives  annually 
run,  or  did  run,  up  to  spawn  in  Campbell's  Pond, 
a  small  body  of  water  that  is  dammed  off  from  the 
harbor,  and  reached  by  a  fishway  only  15  feet  long. 
This  is  the  shortest  alewife  stream  of  which  we 
know. 

In  1896,  when  the  alewife  fishery  was  the  sub- 
ject of  inquiry  by  the  Bureau  of  Fisheries,77  catches 

'•  Such  a  race  has  been  reported  in  Cobbett  Pond,  Rockingham  Co.,  N.  H. 
by  Kendall  (Occ.  Pap.  Boston  Soc.  Nat.  Hist.,  vol.  7,  No.  8, 1908,  p.  3S)  and 
by  Bailey  (Biological  Survey  Merrimac  Watershed,  New  Hampshire  Fish 
and  Oame  Dept.,  1938,  p.  162). 

"  Belding  (Ropt.  Alewife  Fish.  Massachusetts,  Mass.  Dept.  Conserv,  1921) 
has  given  a  very  instructive  report  on  the  alewife  in  Massachusetts. 

"  Smith,  Rept.  U.  S.  Comm.  Fish.  (1898)  1899,  pp.  31-43. 


FISHES  OF  THE  GULF  OF   MAINE 


105 


large  enough  to  be  worth  special  notice  were  re- 
ported from  the  mouths  of  the  St.  Croix,  Dennys, 
Machias,  Medomak,  Penobscot,  St.  George,  Pema- 
quid,  Damariscotta,  and  Kennebec  Rivers;  from 
Casco  Bay;  and  from  sundry  other  shore  localities 
in  Maine;  from  the  Piscataqua  River  system  in 
New  Hampshire ;  from  the  mouth  of  the  Merrimac, 
and  from  Cape  Cod  Bay.  Few  alewives  enter  the 
Merrimac,  now,  so  polluted  is  it,  and  so  obstructed 
by  dams.78  And  Belding  found  them  running  in 
only  about  9  or  10  streams  on  the  Gulf  of  Maine 
coast  of  Massachusetts  in  1920,  out  of  27  streams 
there  that  had  formerly  supported  considerable 
alewife  fisheries.7* 

At  present,  we  learn  from  John  B.  Burns,  of  the 
Massachusetts  Division  of  Marine  Fisheries,  only 
a  few  alewives  manage  to  run  up  the  Merrimac  past 
the  fish  ladder  at  Lowell;  there  is  a  small  but 
regular  run  in  the  Parker  River;  a  few  in  the  Ips- 
wich; a  good  run  in  the  Essex;  a  few  in  the  Saugus; 
perhaps  some  in  Weymouth  Back  River ; M  a  small 
run  in  Wier  River,  Hingham  (really  a  brook);  a 
few  in  Bound  Brook,  Cohasset;  a  large  run  in 
Herring  Brook,  Pembroke  (tributary  to  North 
River)  yielding  about  1,000  barrels  yearly;  in- 
creasing numbers  in  Jones  River,  Kingston,  which 
had  been  restocked  previous  to  1938  when  a  fish 
ladder  was  installed;  several  thousand  run  yearly 
up  Barnstable  Mill  Pond  Brook ;  an  improving  run 
in  Stony  Brook,  Brewster,  where  a  ladder  was  built 
in  1945,  and  a  good  run  in  Herring  River  (really 
only  a  brook)  in  Wellfleet,  Cape  Cod. 

The  first  alewives  ordinarily  appear  early  iD 
April  in  the  few  streams  tributary  to  Massachu- 
setts Bay  that  they  still  frequent,  and  equally 
early  (March  or  April)  in  the  St.  John  River,  New 
Brunswick,  according  to  McKenzie;81  but  their 
date  of  arrival  varies  considerably  from  stream  to 
stream,  according  to  local  conditions.  Thus  few 
are  seen  in  the  streams  of  Maine  until  late  April  or 
early  May;  the  first  alewives  appeared  in  1915,  for 
example,  in  Campbell's  Creek,  Booth  Bay  Harbor, 
on  April  20.  And  the  earliest  good  runs  on  the 
Nova  Scotia  shores  of  the  open  Gulf  and  of  the 
Bay  of  Fundy  may  come  as  early  as  April  (streams 
of  Yarmouth,  Annapolis,  Hants,  and  Colchester 

'*  Fishways  recently  constructed  now  allow  a  few  to  ascend  beyond  Lowell' 
Massachusetts. 

"  See  his  report  on  the  Alewife  Fishery  of  Mass.  (Mass.  Dept.  of  Conser- 
vation, Div.  Fish,  and  Game,  1921)  which  gives  much  information  as  to  the 
status  of  the  alewife  in  Massachusetts  streams. 

"  Stocked  with  28.000  adult  fish  in  1949,  and  fish  ladders  under  construction. 

■'  Eept.  Biol.  Board  Canada  (1931)  1932,  p.  34. 


Counties),  in  May  (Digby  and  King's  County 
streams),  or  not  until  June  (Cumberland  County.82 
Successive  runs  follow  thereafter,  all  around  the 
Gulf,  until  well  into  June,  the  later  runs,  going  up, 
passing  the  earlier  spawners  coming  down.  In 
1915,  we  saw  this  happening  in  Campbell's  Creek, 
Boothbay,  on  May  20.  And  alewives  have  been 
seen,  descending,  as  late  as  August  20,  in  Massa- 
chusetts streams. 

The  extreme  range  of  temperature  within  which 
eggs  are  spawned,  in  Gulf  of  Maine  tributaries,  is 
not  known;  probably  the  bulk  of  production  takes 
place  between  about  55°  and  about  60°. 

Numerical  abundance. — In  1896 M  reported 
catches  were  2,677,972  individual  alewives 
(1,356,755  lb.)  for  Cape  Cod  Bay  and  for  the 
Merrimac  River  combined;  526,500  (293,671  lb.) 
for  New  Hampshire  streams;  and  5,832,900 
(3,388,326  lb.)  from  the  rivers  and  streams  and 
coast  of  Maine.  The  reported  catch  was  5,843,000 
pounds 84  for  the  New  Brunswick  shore  of  the 
Bay  of  Fundy  that  year;  1,609,400  pounds  for  the 
Nova  Scotia  side  and  for  the  west  coast  of  Nova 
Scotia,  or  about  10,510,000  and  about  2,895,000 
individual  fish,  respectively,  assuming  that  the 
average  weight  was  about  the  same  as  that  for 
the  alewives  of  Maine.  We  thus  arrive  at  a  total 
catch  for  the  Gulf  of  Maine  of  something  like  22 
million  individual  fish  at  that  time  and  actually 
somewhat  more,  for  the  canvass  certainly  was  not 
100  percent  complete. 

The  run  was  much  greater  then  in  the  St.  John 
River  system  than  in  any  other  Gulf  of  Maine 
river  and  doubtless  is  still.  The  Damariscotta 
River,  ranking  second,  was  about  one-third  as 
productive  as  the  St.  John;  the  Merrimac,  St. 
George,  and  Penobscot  Rivers  only  something 
like  one-tenth  as  productive  each.  Casco  Bay 
yielded  about  one-sixth  as  many  alewives  as  the 
St.  John  River,  the  shore  line  of  Cape  Cod  Bay 
about  one-fifth  as  many.86  And  the  catch  of  the 
St.  John  River  system  (including  Kennebecasis 
Bay)  still  was  about  five  times  as  great  in  1931  as 
that  for  any  of  the  other  counties  of  New  Bruns- 

»  According  to  McKenzie,  Kept.  Biol.  Board  Canada  (1931)  1932,  p.  34. 

13  A  special  study  of  the  alewife  fishery  was  made  for  that  year,  seo  Smith, 
Eept.  U.  S.  Comm.  of  Fish.   (1896)  1899,  pp.  33^13. 

•*  The  Canadian  catches  for  the  year  were  reported  in  barrels;  the  conver- 
sion factor  used  is  200  pounds  per  barrel. 

**  Reported  catches  for  1896  were  about  4,234,000  pounds  for  the  St.  John 
River  system;  1,390,612  pounds  for  the  Damariscotta  River,  385,804  pounds 
for  the  St.  George  River,  308,844  pounds  for  the  Penobscot,  472,500  pounds 
for  the  Merrimac,  701,287  pounds  for  Casco  Bay,  and  884,255  pounds  for  Cape 
Cod  Bay. 


106 


FISHERY   BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


wick  or  of  Nova  Scotia  that  border  on  the  Bay 
of  Fundy  or  on  the  open  Gulf.88 

The  alewife  population  of  the  Gulf  is  much 
smaller,  today,  than  it  was  half  a  century  ago. 
Thus  the  catch  was  only  about  one-half  as  great 
for  the  Bay  of  Fundy  in  1945  and  1946  87  as  it  had 
been  in  1896,  and  about  one-third  as  great  for 
Maine  (1,224,600  lb.)  while  the  Merrimac  River, 
yielding  472,500  pounds  in  1896,  yielded  less  than 
3,000  pounds  in  1945.88  And  though  alewives 
may  seem  almost  incredibly  numerous  when 
crowding  into  some  stream,  they  made  but  a 
sparse  population,  even  in  their  days  of  greatest 
plenty,  when  spread  over  the  coastal  waters  of 
our  Gulf,  as  compared  to  the  sea  herring. 

Importance. — Alewives  are  excellent  food  fish 
and  they  are  marketed  both  fresh  and  salted,  and 
are  preferred  by  many  to  the  sea  herring.  They 
are  good  bait  for  cod,  haddock,  and  pollock;  and 
their  scales  commanded  a  high  price  for  use  in  the 
manufacture  of  artificial  pearls  for  a  brief  period 
during  the  first  world  war  and  for  a  few  years 
afterward.89  By  far  the  greater  part  of  the  catch 
of  alewives  is  made  in  the  lower  reaches  of  the 
streams  that  they  enter  to  spawn,  in  weirs,  in 
dip  nets  or  in  haul  seines  according  to  locality. 
Most  of  those  taken  in  outside  waters  (as  in 
Casco  and  Cape  Cod  Bays)  are  either  gill  netted 
or  are  picked  up  in  the  fish  traps. 

»  McKenzie,  Kept.  Biol.  Board  Canada  (1931)  1932,  p.  34. 

87  5,051,100  pounds  and  4,517,500  pounds,  respectively. 

M  The  reported  catch  for  Essex  County,  Massachusetts,  in  that  year  was 
2,700  pounds,  only  a  part  of  which  was  from  the  region  of  the  Merrimac. 

*  For  details,  see  Report,  Division  of  Fish  and  Game,  Mass.  (1920)  1921, 
p.  140. 


Blueback  Pomolobus  aestivalis  (Mitchill)  1815 

Glut  herring;  Summer  herring;  Blackbelly; 

Ktack 

Jordan  and  Evermann,  1896-1900,  p.  426. 

Description. — Bluebacks  and  alewives  are  diffi- 
cult to  distinguish;  experienced  fishermen  who 
recognize  the  existence  of  the  two  separate  fish 
cannot  always  tell  them  apart,  so  closely  do  they 
resemble  one  another  in  general  appearance.  The 
most  obvious  external  difference  between  them  is 
that  the  back  of  the  blueback  is  definitely  blue 
green,  that  of  the  alewife  gray  green.  But  this 
applies  only  to  fresh-caught  fish;  preserved  speci- 
mens do  not  differ  much  in  color,  or  fish  that  have 
been  on  ice  for  more  than  a  short  time.  Another 
external  difference  is  that  the  eye  of  the  blueback 
is  only  about  as  broad  as  the  distance  from  front 
of  eye  to  tip  of  snout  (or  slightly  broader),  but  is 
appreciably  broader  than  that  in  the  alewife;  the 
blueback,  too,  with  body  about  3%  times  as  long 
as  deep,  is  a  slightly  more  slender  fish  (on  the 
average)  than  the  alewife,  and  its  fins  are  a  little 
lower,  but  the  two  species  probably  intergrade 
in  both  these  respects. 

The  most  dependable  distinction  between  the 
two  (though  requiring  the  use  of  a  knife)  is  that 
the  lining  of  the  belly  cavity  is  sooty  or  blackish  in 
the  blueback,  but  pearl  gray  or  pinkish  gray  in  the 
alewife.  We  have  yet  to  see  a  specimen  that 
could  not  be  named  as  the  one  or  the  other  on  this 
basis  alone,  unless  so  poorly  preserved  that  the 
original  shade  of  the  cavity  could  no  longer  be 
determined. 


Figure  47. — Blueback  (Pomolobus  aestivalis),  Chesapeake  Bay  region  specimen.     From  Goode.     Drawing  by  H.  L. 

Todd. 


FISHES   OF  THE  GULF  OF   MAINE 


107 


Color. — Dark  blue  or  bluish  gray  above,  the 
sides  and  belly  silvery,  with  coppery  reflections  at 
least  in  some  waters;  lining  of  the  belly  sooty  or 
blackish. 

Size. — The  blueback  attains  about  the  same 
size  as  the  alewife,  i.  e.,  15  inches;  the  adults 
average  about  11  inches  in  length  and  about  7 
ounces  in  weight. 

Habits. — The  blueback,  like  the  alewife,  makes 
its  growth  in  salt  water,  but  runs  up  into  fresh 
water  to  spawn.  And  its  breeding  habits  do  not 
differ  in  any  important  particular  from  those  of  the 
alewife,  except  that  it  "runs"  later  in  the  season, 
does  not  run  up  as  far  above  tidewater,  and  does 
not  spawn  until  the  water  is  much  warmer,  70°  to 
75°  instead  of  55°  to  60°.w  The  eggs,  about  1  mm. 
in  diameter,  sink  like  those  of  the  alewife,  and 
stick  to  anything  they  may  chance  to  touch. 
Incubation  occupies  only  about  50  hours  at  a 
temperature  of  72°.  The  young  are  30  to  50  mm. 
long  within  a  month  and  already  show  most  of  the 
diagnostic  characters  of  the  adult.  Evidently 
they  soon  find  their  way  down  to  the  sea,  for  blue- 
backs  of  50  mm.  have  been  seined  in  abundance  in 
Rhode  Island  waters  late  in  July.91  Nothing 
whatever  is  known  of  their  later  rate  of  growth. 
The  spent  fish,  return  to  sea  shortly  after  spawning 
as  do  alewives.  Practically  nothing  is  known  of 
their  movements  in  the  sea,  except  that  they  are 
schooling  fishes.  The  fact,  however,  that  7  were 
trawled  by  the  Albatross  II  on  March  5,  1931  about 
100  miles  off  Cape  May,  N.  J.,  suggests  that  the 
blueback  moves  out  from  land  and  passes  the  cold 
season  near  the  bottom. 

We  need  only  note  further  that  the  blueback  is 
as  gregarious  as  the  herring  or  alewife;  that  it  is 
equally  a  plankton  feeder,  subsisting  chiefly  on 
copepods  and  pelagic  shrimp,  as  well  as  on  young 
launce  and,  no  doubt,  on  other  small  fish  fry. 

Qeneral  range. — -This  is  a  more  southern  fish  than 
the  alewife,  occurring  along  the  American  coast  as 
far  south  as  northern  Florida;  as  far  north  as 
southern  New  England  in  abundance,  perhaps  less 
regularly  in  the  Gulf  of  Maine  though  widespread 

H  The  early  development  and  larval  stages  of  the  blueback  are  described 
by  Kuntz  and  Radclifle  (Boll.  U.  S.  Bur.  Fish.,  vol.  35,  1918,  pp.  87-134). 

"  In  Chesapeake  Bay,  Hildebrand  and  Schroeder  (Bull.  U.  S.  Bur.  Fish., 
vol.  43,  1928,  p.  88)  found  that  while  most  of  the  young  bluebacks  pass  out  to 
sea  during  the  summer  and  fall,  some  remain  in  the  deeper  holes  over  the  win- 
ter. By  the  following  March  when  about  a  year  old  these  are  about  3^  to 
4  Inches  long;  those  In  the  sea  may  grow  faster  than  this. 


there,  and  known  definitely  as  far  north  as  Cape 
Breton,  Nova  Scotia:  M  it  spends  most  of  its  life  in 
salt  water  but  runs  up  into  fresh  water  to  spawn. 

Occurrence  in  the  Gulf  of  Maine. — Although 
fishermen  have  recognized  the  existence  of  two 
distinct  species  of  alewives  at  least  since  1816,  it 
is  difficult  to  arrive  at  a  just  idea  of  the  status 
and  migrations  of  this  fish  in  our  Gulf,  because 
fish  reported  as  "bluebacks"  at  sea  sometimes 
turn  out  to  be  alewives,  while  the  late  runs  of 
alewives  are  often  referred  to  as  "bluebacks." 
It  seems,  however,  that  schools  of  bluebacks  are 
to  be  expected  anywhere  between  Cape  Sable  and 
Cape  Cod.  Thus  we  have  seen  "gaspereau"  fresh 
caught  at  Yarmouth,  Nova  Scotia,  that  appeared 
to  be  bluebacks.83  Huntsman  had  specimens  from 
St.  John  Harbor  and  Shubenacadie  River;  they 
are  reported,  at  least  by  name,  from  the  St.  Croix 
River;  from  Dennys  River,  Eastport;  Bucksport; 
Casco  Bay;  Small  Point;  Freeport;  and  sundry 
other  localities  along  the  coast  of  Maine,  as  well 
as  from  the  shores  of  Massachusetts  Bay,  including 
Cape  Cod. 

L.  W.  Scattergood  of  the  U.  S.  Fish  and  Wildlife 
Service  has  sent  us  about  40  typical  bluebacks, 
about  3%  to  bVt  inches  (92-124  mm.)  long  taken 
at  Hodgdon  Island,  Sheepscot  River,  Maine,  June 
14,  1951 ;  and  we  once  saw  thousands  of  fish  taken 
from  a  trap  near  Gloucester,  most  of  which  we 
judged  to  be  bluebacks  from  their  color.94  A  few 
fish  were  reported  as  "bluebacks"  from  Georges 
Bank  during  the  investigation  of  1913,  and  while 
there  is  no  way,  now,  of  checking  whether  these 
actually  were  bluebacks  or  alewives,  the  fact  that 
we  saw  10  bluebacks  about  1  foot  long,  trawled 
by  Albatross  III  at  the  45  fathom  line  off  southern 
New  England,  in  mid-May,  1950,95  shows  that  they 
may  spread  as  far  offshore  as  alewives. 

No  definite  information  is  at  hand  as  to  how 
regularly  alewives  run  into  our  Gulf  of  Maine 
streams,  for  spawning;  or  what  streams  they  enter 
at  all. 

No  distinction  is  made,  commercially,  on  our 
coast  between  the  blueback  and  the  more  abundant 
alewife;  it  is  equally  useful  for  bait  and  for  food. 

«  Dr.  A.  H.  Leim  has  sent  us  four  typical  bluebacki  about  12  lnchss  long , 
taken  at  Cape  Breton,  Nova  Scotia,  in  1950. 
"  We  had  no  chance  to  examine  them  critically. 

•<  We  did  not  then  appreciate  the  desirability  of  positive  identification. 
"  Lat.  40°  06';  long.  71°  38'  W. 


108 


FISHERY   BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


Shad  Alosa  sapidissima  (Wilson)  1811  [Approxi- 
mate date] 

JordaD  and  Evermann,  1896-1900,  p.  427. 

Description. — The  shad  is  a  typical  member  of 
the  herring  tribe  in  all  respects  with  soft-rayed 
dorsal  and  anal  fins  of  moderate  size,  the  former 
situated  above  the  ventrals  and  well  forward  of 
the  middle  of  the  body.  It  has  a  deeply  forked 
tail  and  large  scales  that  are  loosened  easily. 
Unlike  the  sea  herring,  the  shad  has  no  teeth  on 
the  roof  of  the  mouth;  adults  have  no  teeth  at 
all,  although  young  shad  have  small  ones  in  the 
jaws  which  may  persist  until  the  fish  is  a  foot  or 
so  long.  The  shad  agrees  with  the  hickory  shad, 
alewife,  and  blueback,  in  its  deep  body  and  sharp 
saw-edged  belly.  But  it  differs  rather  noticeably 
from  the  hickory  shad  in  its  longer  mouth,  with 
upper  jaw  reaching  to  below  the  rear  edge  of 
the  eye,  and  in  the  fact  that  the  tip  of  its  lower 
jaw  is  entirely  enclosed  within  the  tip  of  the  upper 
when  its  mouth  is  closed.  The  most  clear  cut 
character  distinguishing  shad  from  alewife  and 
blueback  is  that  the  upper  outline  of  the  shad's 
lower  jaw  is  very  slightly  concave,  without  a 
sharp  angle,  the  outline  of  theirs  deeply  concave 
with  a  pronounced  angle.  Furthermore  the  lining 
of  the  shad's  belly  is  very  pale. 

Color. — Dark  bluish  or  greenish  above,  white 
and  silvery  low  on  sides  and  on  belly,  with  a 
dusky  spot  close  behind  the  rear  edge  of  the  gill 
cover,  and  usually  with  one  or  two  longitudinal 
rows  of  indistinct  dusky  spots  behind  it. 

Size. — The  shad  is  the  largest  of  the  herrings 
that  regularly  visit  our  Gulf,  growing  to  a  length 
of  2%  feet.     In  the  Bay  of  Fundy,  according  to 


Leim  M  shad  weigh  about  %  pound  at  8  inches; 
about  %  pound  at  12  inches;  about  IK  pounds  at 
15  to  16  inches;  about  2%  pounds  at  about  20 
inches;  and  about  4 %  pounds  at  23  to  24  inches, 
though  with  variations  according  to  their  condi- 
tion. Adult  males  weigh  from  IK  to  6  pounds; 
females  from  3K  to  8  pounds.  Shad  are  occasion- 
ally reported  to  12  pounds,  and  the  older  writers 
mention  shad  of  14  pounds,  but  none  so  large  has 
been  credibly  reported  in  the  Gulf  of  late  years. 

Habits. — The  shad,  like  the  alewife,  spends 
most  of  its  life  at  sea,  and  makes  most  of  its 
growth  there,  but  runs  up  into  fresh  rivers  to 
spawn,  the  spent  fish  soon  returning  to  salt  water, 
and  its  fry  soon  running  down  also.  During  their 
stay  in  the  sea  shad  are  schooling  fish,  often  in 
thousands,  and  they  never  reenter  fresh  water 
until  they  return  to  spawn,  though  they  sometimes 
do  appear  in  brackish  estuaries.  Schools  of  shad 
are  often  seen  at  the  surface  in  spring,  summer, 
and  autumn.  In  winter  they  disappear  from 
sight.  Probably  the  shad  of  the  year  winter  near 
the  mouths  of  their  parent  streams;  the  larger 
sizes  somewhat  farther  out  and  deeper.  The  most 
direct  evidence  as  to  the  depths  to  which  they 
may  descend  is  that  shad  have  been  trawled  at 
about  50  fathoms  off  Nova  Scotia  in  March  (see 
footnote  22,  p.  112),  and  at  26  to  68  fathoms  off 
southern  New  England  in  May  (footnote  23,  p. 
112). 

Food. — The  shad,  like  other  herrings,  is  pri- 
marily a  plankton  feeder.  We  have  found  shad 
taken  in  the  Gulf  of  Maine  in  summer  full  of 
copepods  (chiefly  Calanus),  and  the  stomach  con- 


"Contrib.  Canad.  Biol..  N.  Ser.,  vol.  2,  1924,  p.  245,  fig.  41. 


Figdee  48. — Shad  (Alosa  sapidissima),  Chesapeake  Bay  specimen.     From  Goode.     Drawing  by  H.  L.  Todd. 


FISHES   OF  THE  GULF  OF   MAINE 


109 


tents  of  fish  from  the  Nova  Scotia  Coast  of  the 
Bay  of  Fundy  examined  by  Willey  "  consisted 
chiefly  of  the  copepod  genera  Arcatia  and  Temora 
with  other  smaller  ones,  of  mysid  shrimps  and  of 
the  larval  stages  of  barnacles;  while  Leim  M  found 
that  the  shad  in  the  open  Bay  of  Fundy  feed 
chiefly  on  copepods  and  mysids.  Shad  are  also 
known  to  feed  as  greedily  on  the  pelagic  euphausiid 
shrimps  as  herring  do,  on  fish  eggs,  and  even  on 
bottom  dwelling  amphipods,  showing  that  they 
forage  near  the  ground  at  times. 

Occasionally  they  eat  small  fish,  but  these  are 
only  a  minor  item  in  their  general  diet."  Shad,  it 
appears,  take  little  or  no  food  just  prior  to  spawn- 
ing. But  they  will  often  take  an  artificial  fly,  or  a 
live  minnow  when  running  upstream  to  spawn.1 
During  the  past  few  years,  crowds  of  anglers  have 
caught  many  on  flies  in  the  Connecticut  River, 
and  doubtless  could  in  the  few  Gulf  of  Maine 
streams  to  which  shad  still  repair  (p.l  10) . 

Reproduction  and  growth.2 — The  sexually  mature 
fish  enter  the  streams  in  spring  or  early  summer 
when  the  river  water  has  warmed  to  50°  to  55°. 
Consequently  the  shad  run  correspondingly  later 
in  the  year  passing  from  south  to  north  along  the 
coast,  commencing  in  Georgia  in  January;  in 
March  in  the  waters  tributary  to  Pamlico  and 
Albemarle  Sounds;  in  April  in  the  Potomac;  and 
in  May  and  June  in  northern  streams  generally 
from  the  Delaware  to  Canada.  In  the  Kennebec, 
according  to  Atkins,3  the  first  shad  appear  (or  did) 
late  in  April,  with  the  main  run  in  May  and  June; 
the  first  ripe  females  are  caught  the  last  week  in 
May  and  they  begin  to  spawn  about  June  1,  most 
of  them  doing  so  during  that  month,  a  few  in  July, 
and  possibly  an  occasional  fisb  as  late  as  August. 
Probably  these  dates  applied  equally  to  the  Merri- 
mac  in  the  good  old  days  when  shad  were  plentiful 
there,  but  the  season  is  somewhat  later  in  the  St. 
John,  also  in  the  Shubenacadie  as  might  be  ex- 
pected; i.e.,  from  mid-May  until  the  end  of  June.* 

•»  Contrib.  Canad.  Biol.,  N.  Ser.,  vol.  1, 1923,  p.  310. 

"  Contrib.  Canad.  Biol..  N.  Ser.,  vol.  2,  1824,  p.  231. 

»  Leldy  (Proc.  Acad.  Nat.  Sci.  Philadelphia,  Ser.,  2, 1868,  p.  228)  found  30 
small  sand  eels  {Ammodytes)  In  the  stomach  of  a  shad,  probably  caught  In 
Delaware  Bay. 

<  Bean  (Bull.  60,  Zool.,  vol.  9,  New  York  State  Mus.,  1903,  p.  207)  com- 
mented on  this  long  ago. 

•  Accounts  of  the  breeding  habits  of  the  shad  have  been  given  by  Ryder, 
Kept.  U.  S.  Comm.  Fish.  (1885)  1887,  p.  623;  by  Prince  (Supp.  6,  Rept.  Dept. 
Marine  Fish.  Canada,  Fish.  Branch.  1907,  pp.  95-110;  in  the  Manual  of  Fish 
Culture,  published  by  the  U.  S.  Bur.  of  Fish.,  1887;  and  more  recently  by 
Leim  (Contrib.  Canadian  Biol.  N.  Ser.  vol.  2, 1924,  pp.  184-202). 

»  Fish.  Ind.  U.  S.,  Sect.  5,  vol.  1, 1887,  pp.  683-684. 

«  Leim,  Contrib.  Canad.  Biol.,  N.  Ser.,  vol.  2,  No.  11, 1924,  p.  182. 


In  large  rivers  they  run  far  upstream.  In  the 
St.  John  River,  New  Brunswick,  they  ascend 
about  200  miles  to  the  grand  falls  even  today 
according  to  Leim,  and  they  still  run  up  300  miles 
(or  did  recently)  in  the  Altamaha  in  Georgia;  for 
375  miles  in  the  St.  Johns  River,  Florida.  But 
they  could  run  up  only  about  35  miles  at  present 
in  the  Penobscot,  where  they  formerly  ascended 
some  90  miles,  or  44  miles  (to  Augusta)  in  the 
Kennebec,  which  they  formerly  ascended  108 
miles  (to  Carratunk  Falls),  though  none  enter 
either  of  these  rivers  now,  so  far  as  we  know. 
And  the  dams  at  Lawrence,  only  20-odd  miles  up- 
stream, now  stop  any  stray  shad  that  may  still 
enter  the  Merrimac,  which  they  formerly  as- 
cended for   125  miles  to  Lake  Winnepesaukee.8 

In  the  Shubenacadie,  shad  spawn  mostly  in 
temperatures  higher  than  about  54°,  and  spawning 
is  interrupted  if  the  water  chills  below  that, 
temporarily. 

The  fish  select  sandy  or  pebbly  shallows  for 
spawning  grounds,  and  deposit  their  eggs  mostly 
between  sundown  and  midnight.  Females  pro- 
duce about  30,000  eggs  on  the  average,  though  as 
many  as  156,000  have  been  estimated  in  very  large 
fish.  The  spent  fish,  now  very  emaciated,  begin 
their  return  journey  to  the  sea  immediately  after 
spawning.  In  the  Kennebec  they  were  first  seen 
on  their  way  down  about  June  20  and  constantly 
thereafter  throughout  July;  in  the  St.  John  spent 
fish  are  running  down  in  July  and  August.  Ac- 
cording to  Atkins  they  begin  feeding  before  reach- 
ing salt  water  and  recover  a  good  deal  of  fat 
before  moving  out  to  sea. 

The  eggs  are  transparent,  pale  pink  or  amber, 
and  being  semi-buoyant  and  not  sticky  like  those 
of  other  river  herrings  they  roll  about  on  the 
bottom  with  the  current.  The  eggs  hatch  in  12 
to  15  days  at  52°  (12°  C),  in  6  to  8  days  at  63° 
(17°  C),  which  covers  the  range  characteristic 
of  Maine  and  Bay  of  Fundy  rivers  during  the 
season  of  incubation.  And  Leim  has  made  the 
interesting  discovery  that  larval  development  is 
more  successful  in  brackish  than  in  pure  fresh 
water,  with  about  7.5  parts  of  salt  per  thousand 
as  about  the  most  favorable  salinity. 

The  larvae  are  about  9  to  10  mm.  long  at  the 
time  of  hatching,  growing  to  about  20  mm.,  at  21 


»  Stevenson  (Rept.  U.  S.  Comm.  Fish.,  (1898)  1899,  p.  Ill)  has  given  a  table 
of  the  distances  to  which  shad  ascended  various  rivers  then,  and  formerly 
from  the  Penobscot  in  Maine  to  the  St.  Johns  in  Florida. 


110 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


to  28  days.  Shad  larvae  resemble  alewife  larvae, 
being  extremely  slender  with  the  vent  almost  as 
far  back  as  the  base  of  the  tail.6  The  young  shad 
remain  in  the  rivers  until  fall,  when  they  move 
down  to  salt  water;  they  are  now  1%  to  4%  inches 
long,  resembling  their  parents  in  appearance. 

According  to  Leim's  investigation,  based  on 
scale  studies  and  length  frequencies,  shad  in  the 
upper  Bay  of  Fundy,  average  about  5  to  6  inches 
long  when  one  year  old;  9  to  10  inches  long  at  2 
years;  13  to  14  inches  at  3  years;  15  to  16  inches 
at  4  years;  and  18  to  19  inches  at  5  years.  The 
two  largest  he  examined,  about  24%  inches  (62  cm. 
and  63  cm.)  long,  appeared  to  be  7  and  6  years  old, 
respectively.  They  may  grow  somewhat  faster 
in  the  open  Gulf  of  Maine,  to  judge  from  the 
greater  abundance  of  pelagic  crustaceans  on  which 
they  feed  (p.  109).  Most  of  the  spawning  fish  are 
5  years  old  in  the  Shubenacadie,  and  presumably 
in  other  Gulf  of  Maine  rivers;  the  oldest  8  or  9 
years  old. 

General  range. — Atlantic  coast  of  North  America 
from  the  southeastern  coast  of  Newfoundland,7 
which  shad  have  been  known  to  reach  as  strays, 
and  the  estuary  of  the  St.  Lawrence  River,  where 
there  is  a  considerable  population  of  them,8  to 
the  St.  Johns  River  in  Florida;  also  represented 
in  the  Gulf  of  Mexico  by  a  closely  related  species. 
The  shad  has  been  successfully  introduced  on  the 
Pacific  coast  of  the  United  States.  It  runs  up 
rivers  into  fresh  water  to  spawn. 

Occurrence  in  the  Gulf  of  Maine. — When  the 
first  settlers  arrived  in  New  England  they  found 
seemingly  inexhaustible  multitudes  of  shad  annu- 
ally running  up  all  the  larger  rivers  and  many 
of  the  smaller  streams,  with  the  tributaries  of  the 
Gulf  of  Maine  hardly  less  productive  than  the 
Hudson  or  Delaware.  But  one  stream  after 
another  was  rendered  impassable  by  the  construc- 
tion of  dams  near  the  mouth,  for  shad  cannot  or 
will  not  run  up  through  fishways  that  are  readily 
used  by  alewives.  Indeed,  they  have  been 
practically  wiped  out  in  the  Merrimac  River,  as 
appears  from  the  following  compilation :  8 

•  Leim  (Contr.  Canad.  Biol.,  N.  Ser.,  vol.  2,  No.  11,  1924,  p.  195)  gives  a 
detailed  comparison  of  shad  with  alewife  larvae. 

1  The  most  northerly  record  of  a  shad,  on  which  we  have  chanced,  is  one 
taken  In  Bull's  Bay,  near  St.  Johns,  Newfoundland. 

•  See  Vladykov  (Contr.  Dept.  Fish.,  Quebec,  No.  30,  1950,  pp.  121-135, 
and  Natural.  Canad.,  vol.  77, 1950,  pp.  121-135)  for  a  study  of  the  movements 
of  the  shad  in  the  St.  Lawrence  estuary. 

•  Frcm  Stevenson,  Rept.  U.  S.  Comm.  Fish.   (1898)  1899,  p.  262. 


Number  of  shad  caught,  Number  of  shad  caught. 

Year  reported,  or  estimated         Year  reported,  or  estimated 

1789... 830,000  1888 None 

1805 540,000  1889 18 

1835 365,000  1890-1892 None 

1865 50,000  1893 2,020 

1871-1873     (aver-  1894 2,750 

age) 1,942  1895-  — 94 

1880 2,139  1896 7 

1885 130 

The  Gulf  of  Maine  rivers  to  which  shad  are 
known  to  resort  regularly  today  are  the  Annapolis, 
Petit  Codiac,  Shubenecadie,  and  St.  John,  tribu- 
tary to  the  Bay  of  Fundy;  perhaps  the  St.  Croix; 10 
the  only  Maine  rivers  that  see  regular  run9  of  a 
few  shad  are  the  Nonesuch  and  the  Sheepscot." 

A  few  shad  may  enter  other  Gulf  of  Maine 
streams  in  some  years  if  not  yearly,  and  bright 
spots  in  the  shad  picture  are  that  a  considerable 
number  of  adult  shad  ran  up  the  South  River  in 
Marshfield,  Massachusetts,  on  the  southern  shore 
of  Massachusetts  Bay  in  1950,  and  that  there 
has  been  a  run  of  something  like  2,000  shad  yearly 
in  Mill  Creek,  Sandwich,  Mass.,  for  the  past  four 
years. 12  How  successfully  they  may  have  spawned 
in  either  of  these  streams  is  not  known. 

It  appears  that  most  of  the  shad  hatched  in  the 
rivers  tributary  to  the  Bay  of  Fundy,  and  the 
spent  fish  from  there,  remain  in  or  near  the  estu- 
aries where  they  take  to  salt  water;  and  that  most 
of  the  adults  that  survive  the  strain  of  spawning 
return  to  the  parent  stream  to  spawn  again. 
Thus  it  is  only  in  St.  Marys  Bay,  in  Annapolis 
Basin,  in  Cobequid  Bay  and  Minas  Basin,  in 
Chignecto  Bay  and  at  the  mouth  of  the  St.  John 
as  well  as  for  a  few  miles  westward,  that  large 
Fundian  shad  are  caught  in  any  numbers.13  The 
fact,  on  which  Leim  13a  comments  that  "there  is 
not  a  single  record  of  a  shad  ever  having  been 
taken"  at  Grand  Manan  island,  although  this 
"lies  almost  directly  in  the  path  of  any  body  of 


«  The  St.  Croix  once  had  a  large  run  of  shad.  None  were  seen  there  for  8 
or  9  years  prior  to  1915,  but  they  wero  there  in  some  numbers  in  1915  and 
1816,  according  to  investigations  by  H.  F.  Taylor  of  the  U.  S.  Bureau  of 
Fisheries;  their  present  status  there  is  not  known.  They  have  been  entirely 
extirpated  from  the  Saco,  where  they  were  abundant  formerly,  probably  from 
the  Penobscot  and  Kennebec,  and  certainly  from  the  Merrimac,  as  noted 
above. 

«  Information  from  Dr.  O.  E.  Atkinson,  U.  S.  Fish  and  Wildlife  Service. 

»  Reported  to  us  by  John  B.  Burns  of  the  Massachusetts  Division  of 
Fisheries  and  Game. 

«  Leim  (Contr.  Canad.  Biol.  N.  Ser.,  vol.  2,  No.  11,  1924,  fig.  2)  gives  a 
chart  showing  the  location  of  shad  catches  for  the  Bay  of  Fundy. 

"■  Contr.  Canad.  Biol.  N.  Ser.,  vol.  2.  No.  11,  1924.  p.  173. 


FISHES   OF  THE  GTTLF   OF   MAINE 


111 


fish  going  in  or  out  of  the  Bay  of  Fundy,"  is 
especially  significant  as  emphasizing  the  localiza- 
tion of  the  St.  John  shad  near  the  parent  river. 

The  behavior  of  the  St.  John  river  shad  raises 
an  interesting  question,  as  to  the  source  of  the 
young  fish  that  sometimes  congregate  in  the 
Bays  and  among  the  islands  along  the  coast  of 
Maine  (Casco  Bay  especially),  for  there  seem  to 
be  too  many  of  them,  in  some  years,  to  be  credited 
to  the  small  runs  that  still  frequent  the  rivers  of 
Maine  (unless  runs  may  have  been  overlooked  of 
late  m  other  rivers  there). 

Immature  shad,  up  to  2  to  2 K  pounds  in  weight 
are  observed  more  or  less  commonly  in  Cape  Cod 
Bay  near  Provincetown  in  summer  or  autumn  and 
in  the  inner  parts  of  Massachusetts  Bay  (some- 
times taken  in  the  traps  at  Beverly  or  Manchester), 
and  off  Cape  Ann.14  Spent  shad  up  to  10  pounds 
in  weight  (averaging  about  5  pounds),  are  some- 
times reported  by  fishermen  off  the  coast  of  Maine 
west  of  Penobscot  Bay;  near  the  Isles  of  Shoals; 
off  York  Beach,  and  off  Cape  Ann,  in  summer, 
autumn,  and  even  in  December.16 

The  few  mature  shad  with  ripening  sexual  organs 
that  are  picked  up  by  the  haddock  netters  between 
Cape  Ann  and  Portland  in  April  and  May,  most 
often  about  the  Isles  of  Shoals  and  Boon  Island,18 
probably  are  headed  for  the  rivers  of  Maine. 

Larger  numbers  of  fish  are  seined  in  September 
and  October,  in  the  neighborhood  of  Mount  Desert 
Island,  where  they  have  been  the  object  of  a  frozen 
fish  industry  in  some  years.17  These,  like  the  green 
fish  mentioned  above,  seem  far  too  numerous  to  be 
accounted  for  by  the  small  production  that  still 
takes  place  in  the  rivers  of  Maine.  Some  few  of 
them,  it  seems,  are  Bay  of  Fundy  fish,  for  one  of  a 
batch  tagged  near  Mount  Desert  Rock  in  August 
1947,  was  recaptured  in  Kings  County,  New 
Brunswick  (St.  John  River  system)  the  following 
June,  and  a  second  in  the  Petitcodiac  River  that 
July,  while  a  third,  tagged  farther  west  on  the 
coast  of  Maine  in  August  or  September  1948  was 


11  502  barrels  (about  100,400  lb.)  were  taken  In  one  set  of  mackerel  pounds 
at  Provincetown  in  June  1910;  tbe  traps  picked  up  numbers  of  shad  of  about 
14  Inches  from  June  20  to  July  6,  1921,  at  Magnolia  and  Beverly,  where  the 
catch  was  10,300  pounds  In  1945;  and  14  shad  11  to  15W  inches  long  were 
taken  in  one  set  of  traps  at  Barnstable,  on  Cape  Cod  Bay,  October  3,  1950. 

>• 135,000  pounds  of  these  large  spent  flsh  were  caught  near  Gloucester  in 
the  autumn  of  1915;  125  barrels  of  2-  to  5-pound  shad,  some  spent,  near  Seguln 
Island,  July  19, 1925. 

18  A  series  of  shad  from  that  region,  examined  by  the  late  W.  W.  Welsh  In 
April  and  May  1913,  averaged  5  pounds,  all  with  well-developed  sei  organs. 

"  About  250,000  pounds  were  brought  in  to  the  local  freezers  yearly  in  1913, 
IOH.  and  1915. 


recaptured  in  the  St.  John  River  in  May  1950. 
But  it  seems  established  that  most  of  the  medium- 
sized  shad  and  larger  now  found  in  our  Gulf  are 
immigrants  from  the  south,  growing  and  fattening 
on  the  rich  supply  of  plankton  they  find  there, 
but  returning  to  the  rivers  west  and  south  of  Cape 
Cod  to  spawn. 

Direct  evidence  of  this  is  that  one  tagged  in 
Chesapeake  Bay  was  recaught  at  Race  Point,  at 
the  tip  of  Cape  Cod,  39  days  later;  18  one  also  was 
recaptured  near  Gloucester  and  another  near 
Portland  that  had  been  tagged  in  the  Hudson 
River,  while  3  out  of  1,380  tagged  in  New  York 
Bay  were  recaptured  in  the  Bay  of  Fundy  after 
37  days,  75  days,  and  85  days,  respectively,  and  one 
tagged  off  Fire  Island,  N.  Y.,  was  recaught  at  St. 
John,  New  Brunswick,  after  39  days.19  On  the 
other  hand,  18  shad,  from  a  batch  of  236  that  were 
tagged  near  Mount  Desert  Rock  in  August  1947 
were  recaptured  the  next  spring  scattered  along  in 
different  stream  systems  from  the  Connecticut  to 
the  Altamaha  in  Georgia.  Others,  from  this  same 
batch,  were  recaptured  in  the  Connecticut,  in  the 
Hudson,  on  the  coast  of  New  Jersey,  and  in  the 
Pamlico  River,  N.  C,  during  the  next  two  springs. 
And  three  others,  from  a  batch  of  431  tagged 
farther  west  along  the  coast  of  Maine  in  the  sum- 
mer and  autumn  of  1948,  were  recaptured  in  the 
Hudson  River;  three  in  Chesapeake  Bay,  and  one 
in  the  Pamlico  River,  N.  C.20 

The  shad  that  take  part  in  this  intermigration 
must  winter  somewhere  between  their  northern 
feeding  grounds  whence  they  have  vanished 
wholly  by  mid-autumn,  and  their  southern  breed- 
ing streams  near  which  they  do  not  appear  until 
spring.  But  it  is  not  yet  known  where  they  pass 
the  cold  months,  how  deep  down  they  go,  how  far 
offshore,  or  how  active  they  are  then. 

Still  other  shad  are  known  to  make  very  long 
journeys  that  can  hardly  be  fitted  into  any  regular 
migratory  pattern,  and  from  which  they  may  never 
find  their  way  back.  Thus  one  that  was  tagged 
in  the  lower  St.  Lawrence  River  was  recaught  on 
Brown's  Bank  258  days  later;  a  second,  from  that 
same  batch,  was  recaught  in  Cumberland  basin, 
near  Amherst,  Nova  Scotia,  at  the  head  of  the 
Bay  of  Fundy  after  322  days;  a  third  at  Province- 


'•  Vladykov,  Trans.  Amer.  Fish.  Soc,  vol.  67, 1938.  p.  64. 
»  Information  supplied  by  C.  E.  Atkinson,  U.  S.  Fish  and  Wildlife  Service. 
"Information  supplied  by  E.  H.  Hollis  of  the  U.  S.  Fish  and  Wildlife 
Service. 


112 


FISHERY  BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


town  at  the  tip  of  Cape  Cod,  some  1,200  miles 
away  from  where  it  had  been  tagged  444  days 
previous.21  And  one,  from  a  batch  of  weir-caught 
fish  tagged  on  the  coast  of  Maine,  August- 
September,  1948,  was  recaught  in  the  Medway 
River,  outer  coast  of  Nova  Scotia,  a  second,  in  the 
Miramichi  River,  tributary  to  the  southern  side  of 
the  Gulf  of  St.  Lawrence  in  1950,  and  a  third,  off 
Tor  Bay,  eastern  Nova  Scotia  in  1951. 

To  what  extent  the  seasonal  journeys  of  the  shad 
are  passive  with  the  dominant  circulatory  move- 
ments of  the  water,  and  to  what  extent  (if  any) 
they  are  self-directed  is  perhaps  the  most  interest- 
ing question  that  now  faces  us  in  our  studies  of  the 
shad  of  the  Gulf  of  Maine. 

Shad  have  been  trawled  50  to  60  miles  out,  off 
eastern  Nova  Scotia;22  have  often  been  reported 
40  to  50  miles  out  off  the  coast  of  Maine;  also 
25  to  90  miles  out,  off  southern  New  England,23 
and  we  saw  one  trawled  by  the  Eugene  H  in  late 
June,  1951,  on  the  southern  part  of  Georges 
Bank  (lat.  40°52'N.,  long.  67°40'W.),  about  110 
miles  from  the  nearest  land.  Evidently  they  may 
wander  as  far  offshore  as  alewives  do;  perhaps 
even  as  far  as  herring. 

Shad  reared  in  different  regions  may,  perhaps, 
prove  to  differ  enough  in  racial  characters  for 
recognition  when  taken  at  sea,  but  this  is  a  ques- 
tion for  the  future.2* 

Abundance. — The  stock  of  shad  in  the  Gulf  is 
but  a  shadow  in  comparison  with  that  of  colonial 
days. 

In  1896,  the  only  year  for  which  detailed 
information  is  available  as  to  the  numbers  taken 
in  different  streams,  290,122  shad  were  reported 
as  caught  in  the  Kennebec  system,  9,000  in  the 
Pleasant  River,  about  3,000  in  the  Harrington 
River,  only  114  in  the  Penobscot  and  12  in  the  St. 
Croix;  100  in  the  Piscataqua  and  7  in  the  Merri- 


51  See  Vladykov,  Nat.  Canad.,  vol.  77,  1950,  p.  121,  for  a  detailed  account 
of  his  tagging  experiments  on  St.  Lawrence  River  shad. 

*>  Vladykov,  Copeia,  1936,  No.  2,  p.  168,  reports  bet-ween  25  and  30  shad  of 
4-6  pounds,  taken  per  haul,  by  otter  trawlers  in  March,  1035,  southwest  of 
Middle  Ground,  about  lat.  44*25'  N„  long.  61°05'  W..  at  about  50  fathoms. 

n  Two  shad  were  trawled  by  Albatross  ///on  the  eastern  part  of  Nantucket 
Shoals  at  68  fathoms,  and  46  others  at  9  stations  distributed  thence  westward 
to  the  offing  to  Montauk  Point  (long.  71°52'  W.)  at  26-64  fathoms,  May  11-18, 
1950. 

It  w  Vladykov  and  Wallace  (Trans.  Amer.  Fish.  Soo.,  vol.  67,  1937-1938,  pp. 
52-66)  believe  that  Sbubenacadie,  Delaware  River  and  Chesapeake  Bay 
shad  differ  significantly  in  average  number  of  vertebrae,  of  mid  ventral 
scales  and  of  pectoral  fln  rays.  But  Warfel  and  Olsen  (Copeia,  1947,  pp. 
177-183)  doubt  whether  any  distinction  can  be  drawn  between  shad  in  various 
streams  along  our  North  Atlantic  coast,  at  least  as  far  as  average  number  of 
vertebrae  goes. 


mac.25  In  that  same  year  the  catch  was  about 
1,059,000  pounds  for  the  Nova  Scotia  shore  of 
the  open  Gulf  and  for  the  Bay  of  Fundy ; x 
1,404,477  pounds  for  the  rivers  and  coast  of 
Maine;  about  122,932  pounds  (32,782  fish)  for 
the  Gulf  of  Maine  coast  of  Massachusetts,  or  a 
total  of  about  2,586,400  pounds  for  the  Gulf  as 
a  whole.  With  shad  averaging  about  3%  pounds 
in  weight,27  this  corresponds  to  about  690,000 
fish. 

But  the  yearly  catch  was  only  about  one-third 
as  great  for  the  period  1916-1919  as  it  had  been 
in  1896,  whether  for  the  United  States  shores  of 
the  Gulf  or  for  the  Canadian.28  And  it  was  of 
about  that  same  order  of  magnitude  in  1931, 
i.  e.,  677,540  pounds  for  the  Gulf  as  a  whole 
(157,763  pounds  for  Maine,  147,277  pounds  for 
Massachusetts,  237,200  pounds  for  the  Bay  of 
Fundy  and  West  Nova  Scotia  region).  Since 
that  time,  the  catches  have  ranged  between 
10,400  pounds  and  306,000  pounds  for  the  Massa- 
chusetts coast  of  the  Gulf  and  between  9,300 
pounds  and  1,106,800  pounds  for  Maine,  a 
fluctuation  so  extreme  (no  regional  correlation 
appearing)  as  to  suggest  that  market  conditions 
were  the  chief  governing  factor.  On  the  other 
hand  the  catches  for  the  Canadian  shores  of  the 
Gulf  increased  rather  consistently  from  1931  to 
a  total  of  1,287,600  pounds  in  1939  then  declined 
to  around  780,000  pounds  for  1944  and  1946,  a 
rise  and  fall  regular  enough  to  suggest  a  corre- 
sponding fluctuation  in  the  actual  abundance  of 
the  shad.  The  average  yearly  catch  for  the  period 
1944-1946  combined,  was  about  20,000  pounds  for 
Massachusetts,  about  224,050  pounds  for  Maine, 
and  about  780,000  pounds  for  the  Bay  of  Fundy 
and  western  Nova  Scotia. 

Thread  herring  Opisthonema  oglinum  (LeSueur) 
1817 

Jordan  and  Evermann,  1896-1900,  p.  432. 

Description. — The  thread  herring  is  distinguish- 
able at  a  glance  from  all  the  herrings  that  regularly 
inhabit  the  Gulf  of  Maine  by  the  prolonged  last 
ray  (usually  about  as  long  as  the  body  is  deep)  of 
its  dorsal  fin.     It  resembles  the  gizzard  shad  of 

■'  Stevenson,  Rept.  U.  S.  Comm.  Fish  (1898)  1899  pp.  265-269. 

18  These  catches  were  reported  as  "barrels"  presumably  of  200  pounds  each. 

"  Stevenson,  Rept.  U.  S.  Comm.  Fish  (1898)  1S99,  p.  121. 

« About  460,000  pounds  for  the  United  States  coast  of  the  Gulf  and 
about  374,000  pounds  for  the  Bay  of  Fundy  and  in  western  Nova  Scotia 
combined  in  1916-17. 


FISHES   OF  THE  GULF  OF   MAINE 


113 


Figure  49. — Thread  herring  (Opisthonema  oglinum).     Drawing  by  Louella  E.  Cable. 


fresh  and  brackish  waters  farther  south  in  this 
respect,  but  the  two  differ  rather  conspicuously  in 
various  details.  In  the  thread  herring,  the  upper 
edge  of  the  tail  fin  is  about  1  }i  times  as  long  as  the 
head  (only  about  as  long  as  the  head  in  the 
gizzard  shad);  the  point  of  origin  of  the  dorsal  fin 
is  a  little  in  front  of  the  origin  of  the  ventral  fins 
(a  little  behind  in  the  gizzard  shad);  the  distance 
from  the  origin  of  the  ventrals  to  the  origin  of  the 
anal  fin  is  at  least  1  %  times  as  long  as  the  base  of 
the  anal  (only  about  %  to  %  in  the  gizzard  shad) ; 
and  the  anal  fin  is  very  low,  with  its  first  few  rays 
a  little  shorter  than  the  eye  (about  1%  times  as 
long  as  the  eye  in  the  "gizzard").  There  is  no 
danger  of  confusing  a  thread  herring  with  a  young 
tarpon  with  which  it  shares  the  prolonged  dorsal 
ray,  for  its  dorsal  fin  originates  in  front  of  the 
ventrals,  while  the  two  fish  are  far  apart  in  general 
appearance.  This  is  a  rather  thin  fish,  its  body 
about  2}i  to  3  times  as  long  (to  the  base  of  the  tail) 
as  deep ;  the  belly  is  sharp  and  saw  edged ;  the  tail 
deeply  forked  as  in  our  other  herrings.  There  are 
18  to  19  rays  in  the  dorsal  fin,  22  to  24  in  the  anal. 

Color. — Bluish  above,  silvery  on  sides  and  belly. 
The  scales  along  the  back  have  dark  centers,  form- 
ing longitudinal  streaks,  and  there  is  a  faint  dark 
spot  just  behind  the  upper  margin  of  the  gill 
cover;  the  dorsal  and  caudal  fins  have  black  tips. 

Size.- — Maximum  length  about  12  inches. 

General  range. — Atlantic  coast  of  America  in 
tropical  and  subtropical  latitudes,  south  to 
Brazd,  straying  northward  to  Chesapeake  Bay, 
and  occasionally  as  far  as  southern  Massachusetts. 

Occurrence  in  the  Gvlj  of  Maine-. — A  thread 
herring  is  caught  off  southern  New  England 
occasionally;  they  were  even  reported  as  rather 


common  in  Buzzards  Bay  and  in  Vineyard  Sound 
during  the  summer  of  1885.  But  there  is  only  one 
record  of  it  within  the  Gulf  of  Maine,  a  single 
specimen  7  inches  long,  taken  off  Monomoy  Point, 
at  the  southern  angle  of  Cape  Cod,  in  August 
1931. 29  Being  a  tropical  fish,  it  is  not  apt  to 
reach  the  Gulf  except  as  the  rarest  of  strays. 

Menhaden  Brevoortia  tyrannus  (Latrobe)  1802 
Pogy;  Mossbtjnker;  Fat  back 

Jordan  and  Evermann,  1896-1900,  p.  433. 

Description. — This  fish  is  universally  called 
"pogy"  in  the  Gulf  of  Maine  but  no  less  than  30 
common  names  are  in  use  south  of  Cape  Cod.  It 
is  flattened  sidewise  like  all  our  other  herrings, 
has  a  sharp-edged  belly,  and  is  as  deep  proportion- 
ally as  the  shad  (body  about  3  times  as  deep  as 
long),  though  the  general  form  is  altered  when  the 
fish  are  fat.  The  very  large  scaleless  head,  which 
occupies  nearly  one-third  of  the  total  length  of  the 
body,  gives  the  menhaden  an  appearance  so  dis- 
tinctive that  it  is  not  apt  to  be  mistaken  for  any 
other  Gulf  of  Maine  fish.  It  is  likewise  distin- 
guishable from  all  its  local  relatives  by  the  fact  that 
the  rear  margins  of  the  scales  are  nearly  vertical 
(not  rounded),  and  are  edged  with  long  comblike 
teeth  instead  of  being  smooth.  The  dorsal  fin 
originates  over  the  ventrals  or  very  slightly 
behind  them.  We  need  only  point  out  further 
that  the  pogy  is  toothless,  its  tail  deeply  forked, 
its  ventral  fins  very  small,  its  dorsal  and  anal  of 
moderate  size,  its  mouth  large  and  gaping  back  as 
far  as  the  hind  margin  of  the  eye,  and  that  the  tip 
of  its  lower  jaw  projects  beyond  the  upper. 

»  Reported  by  MscCoy,  Bull.  Boston  Soo.  Nat.  Hist.,  No.  61, 1931,  p.  21. 


114 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  50. — Menhaden  (Brevoortia  tyrannus).  A,  egg;  B, 
larva,  newly  hatched,  4.5  mm.;  C,  larva,  23  mm.;  D, 
young  fry,  33  mm.     A-D,  after  Kuntz,  and  Radcliffe. 


Color. — Dark  blue,  green,  blue  gray,  or  blue 
brown  above,  witb  silvery  sides,  belly,  and  fins, 
and  with  a  strong  yellow  or  brassy  luster.  There 
is  a  conspicuous  dusky  spot  on  each  side  close 
behind  the  gill  opening,  with  a  varying  number  of 
smaller  dark  spots  farther  back,  arranged  in 
irregular  rows. 

Size. — Adult  menhaden  average  12  to  15  inches 
in  length,  and  from  two-thirds  to  one  pound  in 
weight.  One  18  inches  long  was  taken  at  Woods 
Hole  in  1876,  and  a  fish  20  inches  long  has  been 
reported.  The  heaviest  of  which  we  have  heard 
was  one  of  1  pound  13  ounces,  taken  at  Orient, 
N.  Y. 

Habits. — The  menhaden,  like  the  herring,  almost 
invariably  travels  in  schools  of  hundreds  or  thou- 
sands of  individuals,  swimming  closely  side  by 
side  and  tier  above  tier.  In  calm  weather  they 
often  come  to  the  surface  where  their  identity 
can  be  recognized  by  the  ripple  they  make,  for 
pogies,  like  herring,  make  a  much  more  compact 
disturbance  than  mackerel  do,  and  "a  much  bluer 
and  heavier  commotion  than  herring,  which  hardly 
make  more  of  a  ripple  than  does  a  light  breeze 
passing  over  the  water,"  as  W.   F.   Clapp  has 


stated  to  us.  Also,  pogies  as  they  fe«d  frequently 
lift  their  snouts  out  of  water,  which  we  have  never 
seen  herring  do,  while  they  break  the  water  with 
their  dorsal  fins,  also  with  their  tails.  And  the 
brassy  hue  of  their  sides  catches  the  eye  (as  we 
have  often  seen),  if  one  rows  close  to  a  school 
in  calm  weather. 

It  is  chiefly  on  warm,  still,  sunny  days  that 
the  menhaden  come  to  the  surface,  sinking  in 
bad  weather;  and  they  are  said  to  come  up  more 
often  on  the  flood  tide  than  on  the  ebb.  It  is 
also  said  (this  we  cannot  vouch  for)  that  the 
fish  work  inshore  on  the  flood  tide  and  offshore 
on  the  ebb. 

Food. — The  menhaden,  formerly  thought  to 
subsist  on  mud,  is  now  known  to  feed  chiefly 
on  microscopic  plants  (particularly  diatoms)  and 
on  the  smallest  Crustacea.30  It  sifts  these  out  of 
the  water  with  a  straining  apparatus  in  the  shape 
of  successive  layers  of  comb-like  gill  rakers  as 
efficient  as  our  finest  tow  nets.  No  other  Gulf 
of  Maine  fish  has  a  filtering  apparatus  comparable 
to  that  of  the  pogy,  nor  has  it  any  rival  in  the 

"  For  a  detailed  account  of  the  food  and  of  the  branchial  sieve  of  the  men- 
haden, see  Peck  (Bull.,  U.  S.  Fish  Comm.,  vol.  13, 1894,  pp.  113-124.  pla.  1-8. 


FISHES   OF  THE  GULF  OF   MAINE 


115 


Gulf  in  its  utilization  of  the  planktonic  vegetable 
pasture.  Menhaden  feed,  as  Peck  described,  by 
swimming  with  the  mouth  open  and  the  gill 
openings  spread.  We  have  often  seen  specimens 
in  the  aquarium  at  Woods  Hole  doing  this.31  And 
we  have  watched  small  ones  in  Chesapeake  Bay, 
swimming  downward  as  they  feed,  then  turning 
upward,  to  break  the  surface  with  their  snouts, 
still  with  open  mouths. 

The  mouth  and  pharyngeal  sieve  act  exactly 
as  a  tow  net,  retaining  whatever  is  large  enough 
to  enmesh,  with  no  voluntary  selection  of  particu- 
lar plankton  units.  The  prey  thus  captured  (as 
appears  from  the  stomach  contents)  includes  small 
annelid  worms,  various  minute  Crustacea,  schizo- 
pod  and  decapod  larvae,  and  rotifers,  but  these 
are  greatly  outnumbered  as  a  rule  by  the  sundry 
unicellular  plants,  particularly  by  diatoms  and 
by  peridinians.  And  the  food  eaten  at  a  given 
locality  parallels  the  general  plankton  content  of 
the  water,  except  that  none  of  the  larger  animals 
appear  in  the  stomachs  of  the  fish  on  the  one 
hand,  nor  the  very  smallest  organisms  (infusoria, 
and  certain  others  such  as  the  coccolithophorids) 
on  the  other.  The  menhaden,  in  short,  parallels 
the  whalebone  whales,  the  basking  shark,  and 
the  giant  devil  rays  in  its  mode  of  feeding,  except 
that  its  diet  is  finer  because  its  filter  is  closer 
meshed. 

Peck  has  calculated  from  observations  on  the 
living  fish  that  an  adult  menhaden  is  capable  of 
filtering  between  6  and  7  gallons  (about  24  to 
28  liters)  of  water  per  minute,  and  while  the 
fish  do  not  feed  continuously  this  will  give  some 
measure  of  the  tremendous  amount  of  water  sifted 
and  of  plankton  required  to  maintain  the  hordes 
in  which  these  fish  congregate.  The  abundance 
of  microscopic  plants  in  the  water  of  bays  and 
estuaries,  and  along  the  coast  has  often  been 
invoked  to  explain  the  concentration  of  menhaden 
close  to  shore. 

Enemies. — No  wonder  the  fat  oily  menhaden, 
swimming  in  schools  of  closely  ranked  individuals, 
helpless  to  protect  itself,  is  the  prey  of  every  pre- 
daceous  animal.  Whales  and  porpoises  devour 
them  in  large  numbers;  sharks  are  often  seen  fol- 
lowing the  pogy  schools;  pollock,  cod,  silver  hake, 
and  swordfish  all  take  their  toll  in  the  Gulf  of 

"  Apparently  Ehrenbaum  (as  quoted  by  Bullen,  Jour.,  Mar.  Biol.  Assoc. 
United  Kingdom,  vol.  9,  1910-13,  pp.  394-403)  was  not  acquainted  with  the 
habits  of  menhaden  when  he  wrote  to  the  effect  that  no  fish  eat  plankton 
Indiscriminately,  or  swim  about  habitually  with  open  mouth  when  feeding. 


Maine,  as  do  weakfish  south  of  Cape  Cod.  Tuna 
also  kill  great  numbers.  But  the  worst  enemy  of 
all  is  the  bluefish,  and  this  is  true  even  in  the  Gulf 
of  Maine  during  periods  when  both  bluefish  and 
menhaden  are  plentiful  there  (p.  384).  Not  only 
do  these  pirates  devour  millions  of  menhaden  every 
summer,  but  they  kill  far  more  than  they  eat. 
Besides  the  toll  taken  by  these  natural  enemies, 
menhaden  often  strand  in  myriads  in  shoal  water, 
either  in  their  attempt  to  escape  their  enemies  or 
for  other  reasons,  to  perish  and  pollute  the  air  for 
weeks  with  the  stench  of  their  decaying  carcasses. 

Breeding  and  growth. — Very  little  is  known  about 
the  breeding  habits  of  the  menhaden,  except  that 
it  spawns  at  sea  and  that  the  chief  production  of 
eggs  takes  place  south  of  our  limits.  According  to 
observations  at  Woods  Hole,32  the  main  body  of 
the  fish  off  southern  New  England  spawn  in  June, 
continuing  through  July  and  August;  even  into 
October  as  in  1915,  when  the  Grampus  collected 
eggs  and  larvae  in  Nantucket  Sound  and  westward 
from  Martha's  Vineyard  in  that  month.  And  re- 
ports of  spent  fish  in  the  Gulf  of  Maine  in  July 
and  August,  with  others  approaching  maturity, 
suggest  that  the  menhaden  is  a  summer  spawner 
there  also.  We  have  found  no  eggs  in  our  tow- 
nettings  north  of  Cape  Cod  (young  fry  were  taken 
in  abundance  in  Casco  Bay  in  October  1900),  prob- 
ably because  our  work  there  was  carried  on  during 
a  series  of  poor  menhaden  seasons.  From  Chesa- 
peake Bay  southward  the  spawning  season  appears 
to  be  late  in  the  autumn,  and  in  early  winter. 

Menhaden  eggs  are  buoyant  and  resemble  those 
of  the  European  pilchard  (Clupea  pilchardus) ,  but 
are  easily  distinguished  from  the  eggs  of  any  other 
Gulf  of  Maine  fish  by  their  large  size  (1.5  to  1.8 
mm.  in  diameter),  broad  perivitelline  space,  small 
oil  globule  (0.15  to  0.17  mm.),  and  very  long  em- 
bryo. Incubation  is  rapid  (less  than  48  hours),  as 
Welsh  found  by  experiment.  The  newly  hatched 
larvae  are  4.5  mm.  in  length,  growing  to  5.7  mm. 
in  4  days  after  hatching.  The  dorsal  and  caudal 
fins  first  become  visible  at  a  length  of  9  mm.;  at 
23  mm  all  the  fins  are  well  developed;  scales  are 
present  at  33  mm.;  and  at  41  mm.  the  fry  show 
most  of  the  characters  of  the  adult,  except  that 
their  eyes  are  much  larger,  proportionately.  The 
youngest  larvae  much  resemble  young  herring,  but 
the  fins  are  formed,  the  tail  becomes  forked,  and 

a  By  Kuntz  and  Radcliffe.  Bull.  U.  8.  Bur.  Fish.,  vol.  35, 1918,  p.  119,  who 
describe  the  eggs  and  larvae. 


116 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


the  body  deepens  at  a  much  smaller  size,  a  men- 
haden of  20  mm.  being  as  far  advanced  in  develop- 
ment as  a  herring  of  35  mm.,  which  makes  it  easy 
to  distinguish  the  older  larvae  of  the  two  fish. 

Welsh  concluded  from  examination  of  great 
numbers  of  fry  and  from  measurements  and  scale 
studies  of  fish  of  various  ages  that  menhaden 
hatched  in  summer  (which  would  apply  to  any 
fry  that  might  be  produced  in  the  Gulf  of  Maine) 
are  2%  to  3%  inches  (6  to  8  cm.)  long  by  their 
first  winter;  and  average  about  6%  inches  (16 
cm.)  by  their  second  winter;  fall-hatched  fish 
are  1%  inches  (3  cm.)  and  about  5  inches  (about 
13  cm.)  long,  in  their  .first  and  second  winters, 
with  every  gradation  between  the  two  depending 
on  the  precise  season  when  the  fish  are  spawned.33 
Apparently  sexual  maturity  is  attained  in  the 
season  following  the  third  winter,  and  a  few  of  the 
older  fish  that  Welsh  examined  showed  as  many 
as  9  to  10  winter  wings  on  their  scales. 

General  range. — Coastal  waters  along  the  At- 
lantic coast  of  America  from  Nova  Scotia  to 
eastern  Florida;  represented  in  the  Gulf  of  Mexico, 
and  southward  to  northern  Argentina,  by  a 
series  of  named  forms  that  differ  from  our  northern 
menhaden  in  ways  that  would  not  be  apparent 
to  any  one  but  to  a  trained  student  of  fishes.34 

Occurrence  in  the  Gulf  of  Maine. — The  Gulf  of 
Maine  is  the  northerly  limit  for  the  menhaden; 
St.  Mary  Bay  on  the  west  coast  of  Nova  Scotia 
is  its  most  easterly  known  outpost.  Prior  to 
about  1850  the  pogy  seems  to  have  been  common 
at  the  mouth  of  the  Bay  of  Fundy;  it  was,  indeed, 
reported  by  Perley  as  far  up  the  bay  as  St.  John, 
and  fishermen  spoke  of  it  as  abundant  near 
Eastport  up  to  1845-1850.  But  it  seems  to  have 
abandoned  Fundian  waters  altogether 35  since 
then  except  for  an  occasional  straggler,  and  very 
few  menhaden  have  been  noticed  east  of  Mount 
Desert  and  Jonesport  of  late  years. 

Perhaps  the  most  interesting  aspect  of  the  oc- 
currence of  the  menhaden  in  the  Gulf  of  Maine  is 
that  it  fluctuates  tremendously  in  abundance  there 
from  year  to  year,  periods  of  great  plenty  al- 
ternating with  periods  of  scarcity  or  entire  absence 

•'  Young  menhaden  that  we  collected  at  Woods  Hole  on  September  23, 
1912,  were  SH  to  4  inches  (91-99  mm.)  long;  others  taken  in  Salt  Pond,  Fal- 
mouth. Mass.,  on  November  24,  1949,  were  4H  to  6  inches  long. 

«  See  Hildebrand  (Smithsonian  Misc.  Coll.,  vol.  107,  1948.  No.  18  for  a 
revision  of  the  genus  Breroortia).  One  named  species,  P.  brericaudala 
Goode  1878,  is  known  only  from  Noank,  Conn.;  we  doubt  its  validity. 

••According  to  Huntsman  (Contr.  Canad.  Biol.,  (1921)  1922,  p.  69)  one 
was  taken  in  St.  John  Harbor  in  August,  1919. 


from    our   waters.     Thus    they    were    extremely 
abundant  off  the  coasts  of  Massachusetts  and 
Maine,  every  summer,  for  some  years  prior  to 
1875,  when  a  considerable  fishery  developed  for 
them  in  Maine.    Very  few,  however,  were  taken 
in  the  Gulf  during  the  cold  summer  of  1877  until 
September  and  October,  when  they  were  reported 
as   about    as    abundant   as    normal;    practically 
none  appeared  north  of  Cape  Cod  in  the  year 
1879;  and  they  were  so  scarce  along  the  coast  of 
Maine  for  the  next  six  years  that  it  caused  com- 
ment when  an  occasional  one  was  caught.     In 
1883,  for  instance,  a  few  were  reported  to  the 
U.  S.  Fish  Commission  though  no  schools  were 
seen  and  many  people  thought  they  had  gone  per- 
manently.    But  they  were  once  more  reported 
abundant  off  Maine  and  Massachusetts  in  1886; 
they  were  so  plentiful  as  far  east  as  Frenchman 
Bay  in   1888  that  the  menhaden  fisheries  were 
revived;  they  were  as  plentiful  in  Maine  waters 
in  1889  as  they  had  ever  been  (more  than   10 
million  pounds  taken  there)  and  they  were  still  so 
numerous  in   1890   that  four  fertilizer  factories 
were  established,  and  nearly  90  million  fish  were 
taken  during  that  season.     But  this  period   of 
abundance  was  short-lived,  less  than  half  as  many 
fish  being  caught  in  Maine  waters  (about  41  mil- 
lion) in  1891  as  the  year  before,  while  few  men- 
haden were  taken  or  seen  north  of  Cape  Cod  in 
1892.     They  were  plentiful  enough,  however,  in 
1894,  for  a  single  steamer  to  seine  about  a  million 
fish  off  the  Kennebec  during  that  summer,  while 
582,131  fish  were  taken  in  Boston  Harbor  in  10 
days'  fishing  during  the  last  half  of  that  August. 
Menhaden  were  scarce  again  in  the  Gulf  during 
the  period  1895-1897  but  abundant  again  in  1898, 
when  about  7  million  pounds  were  taken  along  the 
Maine  coast.     They  were  scarce  in  1902  (Maine 
catch  about  300,000  lb.);  reported  as  abundant 
again  north  of  Cape  Cod,  in  1903,  especially  in 
Boston  Harbor;  rare  north  of  Cape  Cod  from 
1904  to  1921,  when  odd  schools  were  seined  along 
the   Massachusetts   and   Maine   coasts   in   some 
summers,  while  few  or  none  were  seen  in  others. 
They  reappeared,   however,  in  such  abundance 
again  in  the  southwest  part  of  the  Gulf  in  the 
summer  of  1922  that  18  steamers  fished  for  them 
successfully  for  some  weeks  in  Massachusetts  Bay, 
when  upwards  of  1,500,000  pounds  were  landed  by 
the  larger  fishing  vessels,  besides  what  the  small 
boats  brought  in.     And  they  were  so  plentiful  at 


FISHES   OF  THE  GULF  OF   MAINE 


117 


least  as  far  north  as  Boothbay  Harbor,  that  about 
2,500  barrels  were  frozen  there,  though  no  large 
schools  were  reported  east  of  that  point. 

The  appearance  of  menhaden  in  such  abundance 
in  the  Gulf  after  so  many  years'  absence  prompted 
the  Bureau  of  Fisheries  to  send  the  steamer 
Halcyon  to  Massachusetts  Bay  that  August,  and 
her  towings  indicated  the  presence  of  much  greater 
quantities  of  diatoms  than  is  usual  at  that  season, 
evidence  that  the  fish  found  a  better  pasture  in 
Massachusetts  Bay  than  in  any  summer  since 
1912.  But  we  hesitate  to  assert  that  it  was  an 
unusually  rich  food  supply  that  attracted  them 
past  Cape  Cod. 

However  this  may  have  been,  there  were  not 
enough  menhaden  in  the  Gulf  to  be  of  any  com- 
mercial importance  from  the  middle  1920's  to  the 
middle  1940's.  But  so  many  visited  Massachu- 
setts Bay,  in  1946  and  1947  that  local  boards  of 
health  were  forced  to  clean  some  of  the  bathing 
beaches  of  the  fish  that  drifted  ashore  from  schools 
netted  for  lobster  bait.  There  were  a  good  many 
in  Maine  waters  in  1948  (reported  catch  145,000 
pounds);36  more  still  in  1949,  when  more  than 
5,000,000  pounds  were  taken  there;  and  about 
8,000,000  pounds  off  Gloucester,37  and  when  small 
fry,  2-3%  inches  (52-95  mm.)  were  taken  in  the 
Sheepscot  River,  December  5-11,  suggesting  that 
some  had  been  reared  in  the  Gulf  that  year.  But 
this  peak  of  abundance  lasted  no  longer  than  the 
peak  had  in  the  early  1920's,  for  there  seem  to  have 
been  far  fewer  menhaden  in  Maine  waters  in  1950 
than  in  1949,  as  there  certainly  were  in  Massachu- 
setts Bay,  where  we  did  not  chance  to  sight  a 
single  school,  and  very  few  were  reported. 

In  the  years  when  menhaden  come,  they  appear 
in  Massachusetts  Bay  about  mid-May;  off  the 
Maine  coast  during  the  last  half  of  May  or  first 
part  of  June.  They  are  most  abundant  during 
July,  August,  and  early  September,  and  most  of 
them  depart  from  the  coast  of  Maine  by  the  middle 
of  October,  from  the  Massachusetts  Bay  region 
by  early  November;  and  it  is  unusual  to  find  a 
single  menhaden  along  these  shores  after  the 
middle  of  that  month,  although  small  ones  have 
been  taken  in  the  Sheepscot  River  as  late  as  the 
first  third  of  December. 

The  universal  belief  among  fishermen,  that  the 
seasonal  appearances  and  disappearances  of  men- 


«  Reported  by  Scattergood,  and  Trefethen,  Copela,  1961,  pp.  93-94. 
•  Reported  by  Scattergood,  Trefethen,  and  Coffin,  Copela,  1951,  p.  298. 


haden  in  the  Gulf  of  Maine  result  from  a  definite 
migration  from  the  south  around  Cape  Cod  in  the 
spring  and  a  return  journey  in  the  autumn, 
probably  is  well  founded. 

The  brevity  of  the  peaks  of  abundance,  the  fact 
that  they  come  at  such  long  intervals,  and  es- 
pecially the  great  local  scarcity  of  young  fish,  are 
arguments  against  the  possibility  that  menhaden 
are  permanent  inhabitants  of  our  gulf,  though  a 
few  fry  may  be  produced  there  in  favorable 
summers,  as  happened  in  1949  (p.  117). 

Menhaden  are  warm  water  fish,  and  our  studies 
of  the  temperatures  of  the  Gulf  of  Maine  cor- 
roborate earlier  observations  to  the  effect  that 
they  never  appear  in  spring  until  the  coastwise 
water  has  warmed  to  50°  or  more,  or  in  abundance 
until  the  temperature  is  several  degrees  higher, 
which  is  in  accord  with  Bean's  M  experience  that 
menhaden  will  not  survive  in  an  aquarium  if  the 
water  chills  below  50°.  No  doubt,  it  is  the  falling 
temperature  of  autumn  that  forces  the  menhaden 
to  leave  the  coasts  of  northern  New  England. 

In  menhaden  years  the  fish  occur  all  along  the 
shores  of  the  Gulf  of  Maine  from  Cape  Cod  to 
Penobscot  Bay,  even  to  Mount  Desert.  Their 
chief  centers  of  abundance  always  lie  in  Massa- 
chusetts Bay  within  a  mile  or  so  of  land,  partic, 
ularly  off  Barnstable  and  in  the  mouths  of  Boston 
and  Salem  Harbors;  in  Casco  Bay;  and  among  the 
islands,  thence  to  Penobscot  Bay.  But  we  have 
never  heard  of  them  entering  water  that  is  appre- 
ciable brackish,  and  in  some  years  they  may  con- 
gregate as  much  as  40  to  50  miles  offshore,  as 
happened  in  1878,  for  instance.  But  we  have 
heard  no  report  of  menhaden  in  the  central  part  of 
the  Gulf  or  on  the  off  shore  Banks.  The  men- 
haden are  thin  when  they  arrive  on  our  coasts 
in  spring,  but  they  put  on  fat  so  rapidly  that 
while  the  average  yield  of  oil  per  thousand  Gulf  of 
Maine  fish  was  about  12  gallons  for  the  whole 
summer  season  of  1894,  it  rose  to  14^  gallons  for 
Boston  Harbor  fish  in  August,  and  to  16  or  18 
gallons  in  September.  It  is  generally  accepted, 
furthermore,  that  fish  taken  on  the  New  England 
coast,  south  or  north,  always  average  larger  and 
fatter  than  those  caught  farther  south. 

Commercial  importance. — The  menhaden  is  one 
of  the  most  important,  commercially,  of  the  fishes 
of  the  Atlantic  Coast  of  the  United  States,  being 
used  for  the  manufacture  of  oil,  fertilizer  and  fish 

»  Rept.  New  York  State  Mas.,  60,  Zool.  9, 1903,  p.  213. 


118 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


scrap.39  In  1946,  when  the  catch  for  the  Gulf  of 
Maine  was  only  about  20,000  pounds,  the  total 
catch  for  the  Atlantic  and  Gulf  States  was  851,129, 
000  pounds;  the  value  of  the  catch  to  the  fisher- 
men was  $7,439,573;  the  value  of  the  products 
made   from   menhaden   was   $18,196,573.     Con- 


siderable numbers  are  used  locally  on  the  Middle 
Atlantic  coast  for  bait.  But  the  menhaden  is  so 
oily  that  it  is  unlikely  to  become  popular  as  a  food 
fish.  Practically  the  entire  catch  of  menhaden  is 
taken  by  purse  seines  and  in  pound  nets;  they 
never  bite  a  baited  hook. 


THE  ANCHOVIES.     FAMILY  ENGRAULIDAE 


The  anchovies  are  small  herring-like  fishes;  but 
they  are  easily  distinguishable  from  the  herrings 
by  the  fact  that  their  mouths  are  not  only  very 
much  larger  and  gape  much  farther  back,  but  are 
on  the  lower  side  of  the  head,  and  are  overhung  by 
the  upper  jaw,  which  projects  like  a  short  piglike 
snout  in  some  species.  Two  anchovies  are  known 
to  occur  in  the  Gulf  of  Maine;  both  are  stragglers 
from  the  south. 

KEY  TO  GULF  OF  MAINE  ANCHOVIES 

1.     Anal  fin  originates  under  the  front  of  the  dorsal;  the 
silvery  lateral  band  is  diffuse;  24  to  27  anal  fin  rays 

Anchovy,  p.  118. 

Anal  fn  originates  under  the  rear  rays  of  the  dorsal; 
silvery  lateral  band  bright  and  well  defined;  20  or 
21  anal  fin  rays Striped  anchovy,  p.  119. 

Anchovy  Anchoa  mitchilli  (Cuvier  and 

Valenciennes)  1848 

Whitebait 

Jordan  and  Evermann  (Slolephorus  mitchilli),  1896- 
1900,  p.  446. 

Description. — The  only  Gulf  of  Maine  fishes  with 
which  one  might  confuse  an  anchovy  are  young 
herring,  smelt,  or  silversides,  but  it  is  easily  dis- 
tinguished from  the  former  by  the  wide  mouth,  as 
just  noted;  by  its  much  larger  eye;  by  the  relative 
positions  of  the  fins  with  the  dorsal  wholly  behind 

■  For  an  account  of  the  menhaden  industry,  see  Harrison,  Inv.  Rept.  No. 
1,  U.  S.  Bureau  of  Fisheries,  1931. 


the  ventrals  instead  of  over  them  and  with  the 
latter  originating  close  behind  the  tips  of  the 
pectorals  when  these  are  laid  back  against  the 
body;  and  by  its  much  longer  anal  fin.  The  lack 
of  an  adipose  fin  behind  the  dorsal  is  sufficient  to 
separate  anchovy  from  smelt  at  a  glance,  while 
the  silversides  (Menidia)  have  two  dorsal  fins  in- 
stead of  one.  The  anchovy  has  large,  thin, 
easily  detached  scales  and  a  deeply  forked  tail. 
This  species  may  be  distinguished  from  the  striped 
anchovy  by  the  fact  that  its  anal  fin  originates 
under  the  front  of  the  dorsal ;  by  its  more  or  less 
diffuse  lateral  band  of  silver;  by  its  more  numerous 
anal  fin  rays  (24  to  27  contrasted  with  20  or  21  for 
the  striped  anchovy),  and  by  its  relatively  small 
size,  for  it  seldom  exceeds  3  inches  in  length.  The 
body  is  about  4  to  5  times  as  long  as  deep  in  both 
anchovies. 

Color.- — This  is  a  whitish  silvery,  translucent 
little  fish,  its  most  characteristic  marking  being  an 
ill-defined  silvery  band  scarcely  wider  than  the 
pupil  of  the  eye,  running  from  the  gill  opening 
back  to  the  caudal  fin.  There  are  also  many  dark 
dots  on  body  and  fins. 

Size. — Seldom  more  than  3K  inches  long. 

General  range.- — Coast  of  the  United  States 
from  Maine  to  Texas,  chiefly  west  and  south  of 
Cape  Cod. 

10  For  a  recent  review  of  the  American  anchovies  see  Hildebrand,  Bull- 
Bingham  Oceanographic  Coll.,  vol.  8,  art.  2, 1943. 


Figure  51. — Anchovy  (Anchoa  mitchilli). 


FISHES  OF  THE  GULF  OF   MAINE 


119 


Occurrence  in  the  Gulf  of  Maine. — -We  mention 
the  anchovy  because  it  has  been  taken  in  Casco 
Bay  and  at  Provincetown.  It  has  no  real  place 
in  the  Gulf  of  Maine  fauna,  seldom  straying  past 
Cape  Cod,  though  it  is  abundant  about  Woods 
Hole  and  thence  westward  and  southward. 
Stragglers  may  be  expected  most  often  in  the  Gulf 
in  midsummer  for  it  appears  from  May  to  October 
in  southern  New  England  waters.  Sandy  beaches 
and  the  mouths  of  rivers  are  its  chief  resorts.  An 
account  of  its  embryology  and  larval  development 
is  given  by  Kuntz.41 

Striped  anchovy  Anchoa  hepsetus  (Linnaeus)  1758 

Jordan  and  Evermann  (Stolephorus  brownii),  1896-1900, 
p.  443. 

Description. — This  anchovy  resembles  the  pre- 
ceding species  closely,  but  its  anal  fin  is  shorter 
(20  or  21  rays)  and  originates  under  the  last  rays 
of  the  dorsal,  and  it  has  a  very  bright  and  well  de- 
fined silvery  band  along  each  side.  It  is  a  larger 
and  more  robust  fish  than  the  other  anchovy, 
often  more  than  4  inches  long. 

Color. — The  bright  silvery  lateral  band,  already 
mentioned,  is  the  most  prominent  marking  on  this 


fish.  Fresh  specimens  are  pale  gray  and  irides- 
cent, the  upper  surface  of  the  head  with  some 
green  and  yellow;  and  the  back  has  dusky  dots. 
The  dorsal  and  caudal  fins  are  more  or  less  dusky 
on  some  specimens. 

Size. — Commonly  4  to  5  inches  long,  maximum 
length  about  6  inches. 

General  range. — Abundant  from  Chesapeake 
Bay  to  the  West  Indies,  and  south  to  Uruguay; 
north  as  a  stray  to  Maine  and  to  the  outer  coast 
of  Nova  Scotia;  a  a  more  southerly  fish  than  the 
other  anchovy. 

Occurrence  in  the  Gulf  of  Maine. — The  claim  of 
this  species  for  mention  in  the  Gulf  of  Maine  is 
based  on  one  record  off  the  mouth  of  the  Penobscot 
River,  near  Portland,  October  8,  1930.43  One 
specimen  was  saved  and  identified,  and  the  her- 
ring fishermen  who  brought  it  in  stated  that  there 
were  "lots  of  them"  on  that  date.  It  is  not  likely 
that  the  striped  anchovy  is  other  than  a  straggler 
to  the  Gulf,  else  it  would  have  been  found  there 
before  this.  As  it  is  a  gregarious  fish,  nearly 
always  traveling  in  small  schools,  it  is  not  aston- 
ishing that  they  may  be  found  together  in  some 
numbers,  on  occasion,  even  out  of  their  usual 
range. 


Figure  52. — Striped  anchovy  (Anchoa  hepsetus),  Somers  Point,  N.  J.,  specimen  100  mm.  long. 
THE   SALMONS.     FAMILY   SALMONIDAE 


The  salmons  are  soft-rayed  fishes  with  no  spines 
in  any  of  the  fins,  with  the  ventrals  situated  on 
the  abdomen  far  behind  the  pectorals,  and  with  a 
fleshy  rayless  "adipose"  fin  on  the  back  behind 
the  rayed  dorsal  fin.  The  presence  of  this  adipose 
fin,  and  its  situation,  separates  them  from  all 
other  Gulf  of  Maine  fishes  except  for  the  smelt, 
capelin  and  the  argentine,  the  pearlsides  (p.  144), 
and  some  of  the  lantern,  viper,  and  lancet  fish 

u  Bulletin.  U.  8.  Bur.  of  Fish.,  vol.  33,  1915,  p.  13. 


tribes  (p.  141).4*  The  blunt  noses,  stout  bodies, 
and  nearly  square  tails  of  the  salmons  distinguish 
them  at  a  glance  from  the  sharper-nosed,  slender, 
forked-tailed  smelts,  their  large  mouths  and  smaller 
eyes  from  the  argentine;  the  absence  of  lumi- 
nescent   organs    distinguishes    them    from    the 


«  Five  were  taken  in  Bedford  Basin,  Halifax,  Nova  Scotia,  on  September 
29,  1931  (Vladykov,  Proc.  Nova  Scotia  Inst.  Sci.,  vol.  19,  193S,  p.  3). 
«  Kendall,  Bull.  Boston  Soc.  Nat.  Hist.,  No.  58, 1931. 
"  Sundry  other  deep-sea  fishes  have  adipose  fins. 


120 


FISHERY   BULLETIN   OP   THE    FISH   AND  WILDLIFE   SERVICE 


pearlsides,  while  the  lantern,   viper,   and  lancet 
fishes  are  of  different  general  aspect. 

Four  salmons  46  occur  in  the  Gulf  of  Maine,  or 
have  recently,  one  of  which,  the  sea  trout,  resorts 
to  tidal  estuaries  at  the  mouths  of  a  few  of  our 
streams;  a  second  and  a  third — the  humpback 
salmon  and  the  silver  salmon — were  introduced 
from  the  Pacific  coast,  leaving  the  Atlantic  sal- 
mon as  a  characteristic  inhabitant  of  the  open 
waters  of  the  Gulf  of  Maine. 

KEY  TO  GULF  OF  MAINE  SALMONS 

1.  Scales  so  small  that  they  are  hardly  visible;  back  with 

vermiculate  markings;  teeth  on  roof  of  mouth  con- 
fined to  a  group  in  front Brook  trout,  p.  120 

Scales  large  enough  to  be  easily  visible;  back  without 
vermiculate  markings;  a  row  of  teeth  runs  back  along 
the  mid  line  of  the  roof  of  the  mouth 2 

2.  Anal  fin  with  only  8-10  rays Salmon,  p.  121 

Anal  fin  with  12  rays  or  more 3 

3.  Back  and  lower  half  of  tail  fin,  as  well  as  its  upper  half, 

conspicuously     marked     with     large     black     spots 

Humpback  salmon,  p.  131 

Back  with  very  small  black  spots  or  none  at  all;  no 

black  spots  on  lower  half  of  tail  fin.Silver  salmon,  p.  133 

Brook  trout  Salvelinus  jontinalis  (Mitchill)  1815 
Sea  trout;  Salter 

Jordan  and  Evermann,  1896-1900,  p.  506. 

Description. — Although  brook  trout  vary  widely 
in  general  form  in  different  streams,  they  are  usu- 
ally salmon-like  in  shape  when  taken  in  salt  water, 
that  is,  about  one-fourth  as  deep  as  long,  tapering 
gracefully  to  a  small  head.     The  nose  of  a  trout, 

•>  A  specimon  of  one  of  the  whitefishes  (probably  Coreoonus  guadrilatiralis 
Richardson)  was  taken  In  the  mouth  of  the  Sissibou  River,  St.  Mary  Bay, 
Nova  Scotia,  September  1919  [Huntsman,  Contr.  Canad.  Biol.,  (1921)  1922, 
p.  59)  straying  down  from  fresh  water.  Whitensh  have  an  adipose  fin,  like 
the  true  salmons,  but  have  a  very  small  mouth,  and  are  flattened  sidewise, 
and  herring-like  in  appearance,  rather  than  salmon-like. 


however,  is  blunter  than  that  of  a  salmon,  and  its 
head  is  longer  in  proportion,  the  total  length  of 
the  fish  (not  counting  the  caudal  fin)  being  about 
four  and  one-half  times  that  of  the  head,49  while 
its  mouth  (gaping  back  of  the  eye)  is  relatively 
larger.  The  general  arrangement  of  the  fins,  in- 
cluding the  "adipose,"  parallels  that  of  the  salmon, 
but  the  ventral  fins  stand  under  the  middle  of  the 
dorsal,  thus  farther  forward  in  relation  to  the 
latter  than  in  its  larger  relative.  All  the  fins,  too, 
are  relatively  larger,  particularly  the  ventrals; 
as  a  rule  the  anal  has  one  less  ray  in  the  trout 
(usually  8)  than  the  salmon,  but  the  number  of 
dorsal  rays  (about  11)  is  the  same.  The  tail  of 
the  sea  trout  is  less  forked  than  that  of  a  young 
salmon  of  equal  size. 

Examination  of  the  scales  and  of  the  teeth  is  the 
most  positive  means  of  distinguishing  brook  trout 
(in  European  terminology  this  is  a  "charr")  from 
young  salmon,  for  the  teeth  on  the  roof  of  the 
mouth  of  the  trout  are  confined  to  a  cluster  near 
the  front,  instead  of  extending  backward  in  a  row 
along  its  midline  as  in  the  salmon ;  and  the  scales 
of  the  trout  are  so  tiny  as  hardly  to  be  visible 
whereas  those  of  the  salmon  are  large  and  easily 
seen. 

Color. — Trout  living  in  salt  water  almost  wholly 
lack  the  yellow  and  red  tints  so  conspicuous  on 
their  freshwater  relatives.  They  are  steel  blue  or 
bottle  green  on  the  back,  with  cheeks  and  sides 
silvery  like  a  salmon  and  with  a  white  belly.  The 
sides  above  the  lateral  line  are  more  or  less  dotted 
with  pale  yellow  spots,  but  the  dark  vermiculate 
markings  so  characteristic  of  the  fresh-water  brook 
trout  are  rarely  seen  on  the  trunk  of  sea  run  fish, 
though  evident  as  wavy  crossbars  on  the  dorsal 

■  Some  trout  are  longer  headed. 


Figure  53. — Brook  trout  (JSalvelinus  fontinalis) ,  about  15%  inches  long. 


FISHES   OF  THE  GULF  OF   MAINE 


121 


fin  and  on  the  corners  of  the  caudal  fin.  The  sides 
and  flanks  below  the  level  of  the  lateral  line  usually 
are  strewn  with  small  pale  vermillion  dots,  but  the 
ventral  fins  are  often  plain  white;  at  most,  the 
pink  edging  so  conspicuous  in  trout  caught  in 
fresh  water  is  faint  on  fish  in  salt  water. 

General  range. — Eastern  North  America,  north 
to  the  outer  coast  of  Labrador,  west  to  Minnesota, 
and  southward  to  Georgia  along  the  Allegheny 
Mountains. 

Occurrence  in  the  Gulf  of  Maine. — Brook  trout 
are  plentiful  in  many  of  the  river  systems  and 
smaller  streams  that  empty  into  the  Gulf  of  Maine. 
Some  of  the  trout  in  some  of  these  seek  salt  water 
after  the  breeding  season,  to  remain  there  over 
the  winter.  This  applies  particularly  to  the 
brooks  that  flow  through  the  sands  of  Cape  Cod, 
several  of  those  on  its  southern  slope  being  famous 
for  their  sea-trout  fishing.  These,  however,  he 
outside  our  present  province,  and  only  a  couple 
of  small  streams  on  the  Massachusetts  Bay  side 
of  the  Cape  still  support  a  race  of  trout  that  run 
down  to  the  sea  regularly.  One  or  two  small 
brooks  tributary  to  Ipswich  Bay,  and  the  Merriland 
River,  emptying  between  Wells  and  Kennebunk- 
port,  Maine,  are  the  only  places  between  Cape 
Ann  and  Cape  Elizabeth  where  we  have  heard  of 
sea  run  trout. 

We  cannot  say  how  generally  sea  trout  may  now 
exist  in  the  streams  of  eastern  Maine,  but  accord- 
ing to  Evermann  47  trout  once  inhabited  the  tidal 
portions  of  many  of  the  brooks  that  empty  into 
Casco  Bay,  and  they  still  may.  Some  of  good 
size  are  caught  also  in  the  Belfast  River  waters, 
tributary  to  upper  Penobscot  Bay.48  Huntsman 
found  no  definite  evidence  of  trout  in  salt  or 
brackish  water  on  the  New  Brunswick  side  of  the 
Bay  of  Fundy,  but  local  inquiry  has  elicited  the 
information  that  there  are  fish  of  this  habit  in  a 
few  streams  (notably  in  Salmon  River)  on  the 
north  and  west  coasts  of  Nova  Scotia,  where 
many  streams  formerly  held  sea  run  trout  that 
have  been  fished  out  long  since. 

The  "sea  trout"  are  indistinguishable  from  the 
ordinary  brook  trout  anatomically.49  They  are 
simply  fish  that  have  the  habit  of  running  down 
to  salt  water,  and  most  of  the  trout  never  leave 

«I  Rept.  U.  S.  Comm.  Fish.,  (1904)  1905,  p.  105. 

«  Towne,  Striped  Bass  Survey,  Maine  Development  Comm.  and  Dept. 
Sea  and  Shore  Fisheries,  1940,  p.  21. 

••  There  Is  another  species  of  sea  trout  (Salvclinus  alpinus)  in  northern 
Canadian  waters  which  is  very  plentiful  along  the  coast  of  northern  Labrador. 

210941—6 


fresh  water,  even  in  streams  offering  free  access  to 
the  sea,  cold  enough  throughout  their  lengths, 
and  harboring  these  "salters"  (as  they  are  called 
on  Cape  Cod).  All  who  have  given  special  atten- 
tion to  our  sea  trout  are  agreed  on  this.  It  is 
still  an  open  question  whether  the  habit  is  hered- 
itary or  whether  it  is  acquired  independently  by 
each  individual  fish.  We  incline  to  the  first  view, 
chiefly  because  sea  trout  are  slow  in  reestablishing 
themselves  in  any  stream  where  they  have  been 
brought  to  a  low  ebb  by  hard  fishing.  The  trout 
that  follow  this  habit  grow  much  more  rapidly  on 
the  abundant  rations  the  salt  estuaries  provide 
than  do  most  of  their  relatives  that  remain  in  the 
brook.  Sea  fish  weigh  from  1  to  3  pounds  in 
streams  where  few  of  the  fresh-water  trout  exceed 
half  a  pound. 

On  Cape  Cod  the  sea  trout  go  down  to  salt 
water  hi  November  immediately  after  spawning, 
to  winter  there.  They  begin  to  run  again  in 
April,  and  all  of  them  are  in  brackish  or  fresh 
water  by  mid-May.  But  it  is  said  that  they  do 
not  appear  until  later  in  the  Nova  Scotia  streams 
tributary  to  the  Bay  of  Fundy  (we  cannot  vouch 
for  this). 

While  in  salt  water  (at  least  along  Cape  Cod) 
the  trout  feed  chiefly  on  shrimps  or  on  gammarid 
Crustacea,  on  mummichogs  (Fundulus),  and  on 
other  small  fish.  Trout  never  stray  far  from  the 
stream  mouths;  hence  they  have  no  place  w  in 
the  fish  fauna  of  the  open  Gulf. 

Salmon  Salmo  salar  Linnaeus  1758 

Atlantic  salmon;  Sea  salmon;  Silver  salmon; 
Black  salmon;  Parr;  Smolt;  Grilse;  Kelt 

Jordan  and  Evermann,  1896-1900,  p.  486. 

Description. — The  Atlantic  salmon  is  a  graceful 
fish,  about  one-fourth  as  deep  as  long,  deepest 
below  the  dorsal  fin,  whence  it  tapers  toward  both 
head  and  tail;  and  oval  in  cross  section.  Its 
head  is  small  (about  one-fifth,  or  less  of  the  fish's 
length,  not  counting  the  caudal  fin),  its  nose  is 
blunt,  eye  rather  small,  and  its  mouth  gapes  back 
to  below  the  eye.  The  dorsal  fin  (about  11  rays) 
stands  about  midway  between  tip  of  snout 
and  base  of  tail  fin;  the  ventrals  are  under  the  rear 
end  of  the  dorsal.  The  anal  is  similar  in  form  to 
the  dorsal  but  has  only  about  9  rays  (7  to  10  have 

»  Trout  are  taken  about  Woods  Hole,  occasionally,  in  winter. 


122 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  54. — Salmon  (Salmo  salar).     Drawing  by  H.  L.  Todd. 


been  recorded),  whereas  the  humpback  has  14 
anal  rays  or  more.  The  tail  is  only  very  slightly 
emarginate  in  adults,  and  is  almost  square  in 
large  fish,  but  is  more  forked  in  fish  that  have  been 
at  sea  for  not  more  than  one  year  ("smolts"  and 

"grilse"). 

Color. — The  salmon  is  silvery  all  over  while  in 
the  sea,  with  brownish  back  and  with  numerous 
small  black  crosses  and  spots  on  head,  body  (chiefly 
above  the  lateral  line),  and  fins.  The  young  fish 
(or  "parr")  are  conspicuously  marked  with  10  or 
11  dark  crossbars  while  in  fresh  water,  alternating 
with  bright  red  spots,  much  like  young  trout. 
Fish  that  have  been  at  sea  for  only  one  year 
(grilse)  are  marked  with  a  larger  number  of  black 
spots  than  the  older  fish. 

Size. — The  largest  salmon  we  find  mentioned 
was  an  English  fish  of  83  pounds.  None  even 
approaching  this  size  is  recorded  from  our  side  of 
the  Atlantic,  where  a  50-pounder  is  unusual, 
though  fish  of  40  pounds  are  not  uncommon  in 
some  of  the  larger  rivers  emptying  into  the  Gulf 
of  St.  Lawrence.  Very  few  fish  reach  40  pounds 
in  the  Penobscot  or  St.  John  Rivers,  and  30- 
pounders  are  unusual  there,  the  usual  run  being 
10  to  12  pounds.  Taking  one  river  with  another, 
large  and  small,  10  pounds  may  be  set  as  a  fair 
average  of  the  mature  Gulf  of  Maine  fish.  A  2- 
foot  fish  will  weigh  about  6  pounds,  one  of  3  feet, 
16  to  20  pounds,  with  allowance  for  individual  and 
seasonal  variation. 

Remarks. — The  teeth  and  the  scales  afford  the 
most  certain  distinction  between  small  salmon  and 
the  New  England  sea  trout  (p.  120).  In  thesalmon 
the  roof  of  the  mouth  is  armed  both  with  a  cluster 
in  front  and  with  a  row  of  stout  conical  teeth 
running  back  along  the  mid-line,  easily  felt  with 
the  finger,  whereas  the  sea  trout  has  the  forward 


group  only.  The  scales  of  the  salmon  are  so  large 
that  they  are  seen  easily,  whereas  those  of  the 
trout  are  so  minute  that  they  are  hardly  visible. 
Old  salmon  sometimes  lose  the  teeth  on  the  roof 
of  the  mouth,  but  large  size  and  large  scales 
identify  them  at  a  glance. 

It  should  also  be  easy  to  tell  an  Atlantic  salmon 
from  a  humpback  (should  any  of  the  latter  still 
exist  in  our  Gulf)  for  the  black  spots  on  the  upper 
part  of  the  body  of  the  humpback  and  on  its  tail 
fin  are  more  close  set  and  much  larger  and  con- 
spicuous than  the  dark  markings  on  a  salmon.  A 
more  precise  difference  is  that  an  Atlantic  salmon 
never  has  more  than  10  rays  in  its  anal  fin,  whereas 
the  humpback  always  has  at  least  as  many  as  12, 
while  most  of  them  have  13  to  17. 

The  danger  will  be  greater  of  confusing  smallish 
Atlantic  salmon  with  silver  salmon,  if  the  attempts 
now  in  progress  to  establish  the  latter  in  our  Gulf 
should  succeed,  for  the  two  fish  look  much  alike. 
A  reliable  criterion  is,  again,  the  number  of  rays 
in  the  anal  fin,  for  the  silver  salmon  always  has 
as  many  as  13  of  these,  an  Atlantic  salmon  never 
more  than  10. 

Life  history.61— It  is  no  wonder  that  the  life  of 
the  salmon  has  been  the  subject  of  much  scientific 
study  and  that  a  whole  literature  has  grown  up 
about  it.  As  everybody  knows,  the  salmon  lives 
the  greater  part  of  its  life  in  the  sea  and  makes 
most  of  its  growth  there  but  spawns  in  fresh  water. 

The  salmon  are  silvery  and  very  fat  when  they 
enter  fresh-water  on  the  spawning  journey,  but 

"  Huntsman  (Bull.  Biol.  Board  Canada,  21,  1931)  has  published  an  exten- 
sive study  of  the  life  history  of  the  salmon  of  the  Maritime  Provinces  of 
Canada,  from  which  we  have  drawn  freely  in  the  following  account.  See 
also  Huntsman  and  others  (Migration  and  Conserv.  of  Salmon,  Pub.  No.  8, 
Amer.  Assoc.  Adv.  Sci.,  1939)  for  discussions  of  the' movements  of  the  salmon 
in  Canadian  and  Newfoundland  waters;  also  Lindsay  and  Thompson  (Rept. 
Newfoundland  Fish.  Res.  Comm.,  vol.  1,  No.  2,  1932)  for  an  account  of  the 
biology  of  the  salmon  in  the  rivers  and  around  the  coasts  of  Newfoundland . 


FISHES   OF  THE   GULF   OF   MAINE 


123 


they  lose  condition  gradually  as  they  work  up- 
stream, for  they  feed  very  little  in  fresh  water,  if 
at  all;  they  make  no  attempt,  for  example,  to 
capture  the  parr  they  meet.  Most  anglers  believe 
that  they  may  occasionally  snap  up  a  small  fish  or 
other  tempting  morsel.  Many  are  caught  on 
artificial  flies,  while  every  salmon  angler  knows 
that  they  will  sometimes  take  a  hook  baited  with 
angleworms  or  with  prawns.  It  has  been  suggested 
that  salmon  recover  the  feeding  habits  of  their 
youth  to  some  extent  after  they  have  spent  some 
time  in  the  river,  for  they  often  rise  to  floating 
insects.  But  the  stomachs  of  salmon  caught  in 
fresh  water  never  contain  anything  but  a  little 
yellowish  green  fluid.  And  the  fact  that  they 
keep  better  with  bellies  intact  than  if  opened  and 
gutted  suggests  that  the  secretion  of  effective 
digestive  juices  has  ceased. 

The  maturing  salmon  of  both  sexes  lose  their 
silvery  sheen  in  fresh  water  during  the  summer 
months,  to  take  on  a  dull  brownish  or  reddish  hue, 
while  the  belly  suffuses  with  some  tint  of  red, 
large  black  spots  develop,  and  the  male  not  only 
becomes  variously  mottled  and  spotted  with  red 
or  orange,  but  his  jaws  elongate,  the  lower  becom- 
ing so  hooked  that  only  the  tips  come  together. 
His  body  becomes  slab-sided,  his  fins  thicken,  and 
his  skin  is  covered  with  slime,  until  altogether  he 
is  but  a  caricature  of  the  beautiful  silvery  creature 
that  came  in  from  the  sea. 

In  small  streams  salmon  may  spawn  only  a 
short  distance  above  the  head  of  tide;  but  they 
may  run  upstream  for  more  than  200  miles  in 
large  rivers  that  are  not  obstructed,  as  they  do  in 
the  St.  John  system  in  New  Brunswick.  In  Gulf 
of  Maine  rivers  they  spawn  in  October  and  early 
November,  on  sandy  or  gravelly  bottom,  the 
females  smoothing  a  shallow  trough  or  redd  and 
covering  the  eggs  with  gravel. 

As  it  is  with  the  fife  of  the  salmon  in  the  sea 
that  we  are  concerned  here,  the  reader  is  referred 
to  Belding  62  and  to  Kendall  M  for  recent  accounts 
of  the  mating  actions  of  the  males  and  females. 
The  spent  fish,  known  as  "kelts,"  "slinks,"  or 
"black  salmon,"  are  thin,  weak,  and  so  exhausted 
that  many  of  them  die.  Most  of  those  that  survive 
in  small  rivers  drop  down  at  once  to  the  sea  after 
spawning.  But  many  of  them  finger  over  the 
winter  in  large  rivers,   improving  somewhat  in 

«  Trans.  Amer.  Fish.  Soc,  vol.  24,  1934,  p.  211. 

•>  Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1, 1938,  p.  65-88. 


condition  and  becoming  more  silvery,  though  they 
take  little  food.  If  they  survive  the  winter  (which 
many  do  not,  for  spawning  leaves  them  thin  and 
exhausted^  they  drop  downstream  to  salt  water  the 
following  spring.64 

The  large  (6  to  7  mm.)  thick-shelled  eggs  lie 
loose  on  the  bottom  and  develop  so  slowly  in  the 
low  temperature  of  winter  that  hatching  does  not 
take  place  until  late  in  the  following  April  or  early 
in  May.  The  newly  hatched  larvae  are  15  to  18 
mm.  (0.6-0.7-inch)  long,  and  carry  a  very  large 
yolk  sac  for  about  6  weeks,  hiding  among  the 
pebbles  of  the  spawning  bed  and  taking  no  food. 
When  the  yolk  sac  is  absorbed  the  little  fish,  now 
known  as  "parr,"  begin  to  swim  and  feed. 

Parr  live  in  fresh  water  for  longer  or  shorter 
periods  according  to  locality  or  to  other  factors 
not  well  understood.  In  the  St.  John,65  and  in 
the  rivers  of  Minas  Basin,  most  of  them  remain 
for  2  summers  and  2  winters,  running  down  to 
the  sea  the  third  summer.  But  Huntsman  has 
found  that  some  few  stay  in  the  Fundian  rivers 
for  3  years.  Most  of  the  salmon  of  the  Penobscot 
spend  2  years  as  parr,  a  few  3  years,  according 
to  Kendall.  It  is  even  possible  that  some  may 
linger  in  Gulf  of  Maine  rivers  for  4,  5,  or  even  6 
years,  as  is  known  to  happen  in  Norway.  And 
Dr.  Huntsman  informs  us  that  some  of  the  male 
parr  in  the  rivers  of  the  Chignecto  Peninsula 
become  sexually  mature  before  visiting  the  sea. 

Parr  may  be  moving  downstream  any  time 
from  late  spring  to  autumn,  but  most  of  them 
probably  make  the  journey  in  June  and  July  in 
Gulf  of  Maine  streams,  when  they  are  5  to  6  inches 
long.  They  put  off  their  barred  and  spotted 
pattern  as  they  near  tidewater,  to  assume  the 
silvery  coat  worn  by  the  salmon  during  his  sojourn 
in  the  sea.     They  are  now  known  as  "smolts." 

Salmon,  small  or  large,  are  voracious  while  in 
salt  water,  feeding  altogether  on  live  bait,  chiefly 
on  fish  and  on  crustaceans.  Among  fishes  avail- 
able to  them  in  this  side  of  the  Atlantic,  launce, 
herring,  alewives,  smelt,  capelin,  small  mackerel, 
haddock,  small  sculpins,  and  even  flatfish  have 
all  been  reported  as  entering  into  their  diet  in  one 
place  or  another.  Salmon  caught  off  Norway  are 
sometimes  packed   full  of  herring,  and  a  hook 

**  They  are  voracious  now,  and  fly-fishing  for  these  "black  salmon"  as 
they  are  called,  Is  a  favorite  sport  nowadays,  especially  In  Mlramlchi  waters 
tributary  to  the  Gulf  of  St.  Lawrence. 

"  Huntsman,  Bull  21,  Biol.  Bd.  Canada,  1931,  p.  31,  based  on  studies  by 
Kerr  and  by  Blair. 


124 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


and  line  fishery  is  carried  on  for  salmon  in  the 
Baltic,  with  herring  for  bait,  while  in  British 
waters  salmon  are  sometimes  caught  on  hooks 
baited  with  launce  and  with  pieces  of  mackerel. 
Launce  and  capelin  had  been  the  chief  diet  of 
thousands  of  salmon  opened  by  Comeau66  in  the 
northern  side  of  the  Gulf  of  St.  Lawrence.  And 
it  is  probable  that  the  salmon  of  the  Bay  of  Fundy 
and  open  Gulf  of  Maine  feed  chiefly  on  herring 
(herring  up  to  5  inches  long  have  been  found  in 
salmon  stomachs  near  Eastport)  and  on  launce, 
taking  alewives  or  any  other  small  fish  as  occasion 
offers,  including  smelts  and  mummichogs  (Fundu- 
lus),  when  they  first  enter  the  estuaries.67 

Salmon  also  feed  greedily  on  euphausiid  shrimps 
(fish  entering  the  Penobscot  have  been  found  full 
of  "shrimp,"  probably  euphausiids) ;  to  some 
extent  on  pelagic  amphipods  (Euthemisto) ,  while 
sand  fleas  (gammarid  crustacean)  are  described 
as  ranking  with  launce  and  herring  as  salmon 
food  in  the  North  and  Baltic  Seas.  Salmon  are 
also  credited  with  eating  crabs.68 

Smolts,  on  the  other  hand,  fall  prey  to  any 
large  predaceous  fish  (they  have  been  found  in 
the  stomachs  of  pollock),  but  salmon  are  so  heavy 
and  strong  after  one  or  two  years'  sojourn  in  salt 
water  that  only  fish  as  large  as  tuna,  swordfish, 
or  the  larger  sharks  can  menace  them.  Their 
worst  enemy  is  the  harbor  seal,  which  is  a  com- 
mon inhabitant  of  the  northeastern  coasts  of  the 
Gulf  of  Maine  and  of  the  Bay  of  Fundy. 

The  young  smolts  grow  so  rapidly  on  the  abun- 
dant diet  the  sea  affords  that  they  usually  reach 
a  length  of  at  least  16  inches  and  a  weight  of  any- 
where from  %  to  7  pounds  after  one  year  at  sea. 
They  are  now  known  as  "grilse."  And  older  sal- 
mon continue  to  put  on  length  and  weight  very 
fast,  as  long  as  they  remain  in  salt  water.  Thus, 
several  St.  John  fish  which  were  tagged  and  re- 
leased in  the  river  in  the  autumn  after  spawning 
and  which  were  recaptured  the  following  summer 
after  wintering  in  the  sea  had  gained  2  to  8  pounds 
in  weight,  one  of  them  more  than  6  inches  in  length. 
Others  which  spent  two  uninterrupted  years  in 
the  sea  (as  shown  by  their  scales)  averaged  about 


*  Life  and  Sport  on  the  North  Shore,  1909,  Quebec. 

i  Kendall  (Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1,  1935,  p.  34)  found 
smelts  in  Penobscot  salmon,  alewives  in  salmon  from  the  St.  John. 

»  See  Kendall  (Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1, 1935,  pp.  33-34) 
for  a  recent  survey  of  the  diet  of  salmon  in  general;  the  Oulf  of  Maine  fish  in 
particular,  with  references.  Etchelbaum  (Cons.  Perm.  Internal.  Explor. 
Mer,  Rapports  et  Proc.  Verb.,  vol.  21,  1916,  p.  84)  records  the  contents  of 
many  salmon  from  the  Baltic  and  from  the  North  Sea. 


10  pounds  heavier  and  6  inches  longer  when  re- 
captured.69 But  they  grow  much  less  rapidly  in 
winter  than  in  summer.  And  they  hardly  grow 
at  all  during  the  years  when  they  spawn  if  they 
enter  the  river  early  in  season,  though  they  con- 
tinue growing  until  later  if  they  enter  late.  Hence 
the  size  of  a  salmon  depends  more  on  the  number 
of  times  it  has  spawned  and  on  the  date  when  it 
enters  its  river  than  on  its  age. 

Most  of  the  exceptionally  large  fish  of  40  to  50 
pounds  are  virgin  females  entering  fresh  water  for 
the  first  time,  but  some  are  fish  that  have  already 
spawned  once.  An  interesting  case  is  that  of  a 
45-pound  2-ounce  fish,  caught  in  the  Moisie  River, 
on  the  north  shore  of  the  Gulf  of  St.  Lawrence, 
June  1950,  by  E.  E.  Steedman,  the  life  history  of 
which  had  been  as  follows:60  hatched  spring  1942; 
went  to  sea  June  1945;  returned  to  river  and 
spawned  there  in  1948;  returned  to  sea  autumn 
1948;  remained  there  until  June  1950;  then  re- 
turned to  the  river,  to  be  caught  on  a  "Lady  Am- 
herst" fly;  age  8  years. 

Some  salmon  become  "river  mature"  and  return 
to  spawn  after  only  one  year  at  sea;  these,  known 
as  grilse,  are  distinguishable  from  the  older  fish 
by  more  forked  tail,  more  slender  body,  thinner 
scales,  and  more  numerous  spots  that  are  blue 
rather  than  black.61  Some  spawn  2  or  3  years  in 
succession,  and  hence  never  grow  large;  others 
spawn  twice  in  alternate  years;  a  few  three  times, 
very  few  oftener.  It  follows  from  this  that  large 
salmon  are  to  be  found  in  the  sea  throughout  the 
year,  though  fewer  of  them  in  summer  when  the 
spawning  fish  are  in  the  rivers,  than  in  winter 
when  the  whole  stock  is  in  salt  water  except  for 
the  "parr,"  a  few  immature  grilse  (p.  129),  and 
such  of  the  spent  fish  as  winter  in  the  rivers. 
Some  spawn  only  once  after  3,  4,  or  even  5  years 
at  sea,  growing  to  a  great  size  meantime.  But 
very  few  salmon  five  to  be  more  than  8  or  9 
years  old,  including  the  time  spent  in  fresh  water 
as  parr. 

Our  ignorance  of  the  way  of  life  of  the  salmon 
in  the  sea  has  recently  been  characterized  as 
abyssmal.  Certainly  they  are  swift  swimmers, 
and  the  nature  of  the  catches  suggests  that  they 

»  Huntsman  (Bull.  Biol.  Board  Canada,  No.  21, 1931)  gives  an  interesting 
account  of  these  tagging  experiments,  from  which  this  summary  is  drawn. 

»  As  worked  out  from  its  scales  by  Dr.  D.  L.  Belding,  and  reported  in 
Field  and  Stream,  August  1951,  p.  10. 

•'  It  is  commonly  stated  that  this  applies  chiefly  to  the  males.  But  Hunts- 
man (Bull.  Biol.  Board  Canada,  No.  21, 1931,  pp.  18-19)  has  found  that  grilse 
of  both  sexes  spawn  in  the  small  rivers  at  the  head  of  the  Bay  of  Fundy. 


FISHES   OF  THE  GULF   OF   MAINE 


125 


live  scattered  for  the  most  part.  But  at  least 
one  case  has  come  to  our  notice  of  a  school  seen, 
and  some  of  them  netted.62  While  salmon  often 
leap  in  the  esturaries  on  their  return  journey 
and  in  the  rivers,  we  have  never  heard  of  one 
doing  so  at  sea.  And  they  keep  so  constantly 
to  the  mid-depths  that  they  are  seldom  seen  at 
the  surface,  except  in  the  estuaries.  But  this 
rule  has  its  exceptions,  for  the  school  mentioned 
above  was  sighted  at  the  surface,  where  they  were 
mistaken  for  pollock.  On  the  other  hand,  there 
is  no  reason  to  suppose  that  many  of  the  Gulf  of 
Maine  salmon  descend  to  any  great  depth,  winter 
or  summer.  The  weirs,  gill  nets,  and  other  gear 
that  yield  so  many  in  various  regions,  are  all 
operated  in  rather  shoal  water  (the  Baltic  hook- 
and-line-fishery  is  carried  on  at  about  1  %  fathoms) . 
Dr.  Huntsman  informs  us  that  salmon  are  taken 
on  hand  lines  in  mid-winter  in  the  Bay  of  Fundy. 
They  are  caught  occasionally  on  long  lines  in  the 
Gulf,  and  otter  trawlers  get  stray  salmon  on  the 
offshore  Banks  (p.  126),  proof  that  at  least  some 
may  go  as  deep  as  50  fathoms  or  so,  while  diet 
(p.  124)  proves  that  they  sometimes  feed  near 
bottom  if  not  actually  on  it. 

General  range. — Coastal  waters  of  both  sides 
of  the  North  Atlantic,  entering  rivers  to  spawn. 
On  the  European  side  its  range  extends  northward 
well  within  the  Arctic  Circle;  southward  to  the 
Mifio  River,  at  the  boundary  between  Spain 
and  Northern  Portugal,  perhaps  with  a  few 
reaching  the  Duero  River,  midway  of  Portugal.63 
It  occurs  in  a  few  rivers  in  western  Greenland.64 
On  the  American  side  salmon  ran  up  all  suitable 
rivers,  formerly,  from  northeastern  Labrador 
to  the  Housatonic  emptying  into  Long  Island 
Sound;  perhaps  the  Hudson  also.  The  northern 
limit  of  the  commercial  fishery  for  it  on  the 
American  side  is  only  about  latitude  54°  N. 
(Indian  Harbor,  north  shore  of  Hamilton  Inlet). 
And  while  it  is  known  to  range  to  Hudson  Strait,66 
reports  of  it  from  stream  mouths  northward  from 
Hamilton  Inlet  seem  often  to  have  been  based 


*  Kendall.  Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1,  1935,  p.  32. 

"  This  is  the  southern  European  limit  given  by  Berg  (Zoogeographies. 
vol.  1,  Pt.  2, 1932,  p.  112. 

"  Jensen,  Fauna  of  Greenland,  vol.  1,  Pt.  3,  Fishes,  192S,  pp.  3  and  4,  Copen- 
hagen. 

"  Vladykov  (Contrib.  Canad.  Biol.,  N.  Sor.,  vol.  8,  No.  2,  1933,  p.  18, 
fig.  1)  shows  a  locality  record  near  Fort  Chimo,  and  there  are  salmon  In  the 
rivers  of  the  eastern  part  of  Ungava  Bay. 


on  the  sea  run  form  of  the  Arctic  charr  Salvelinus 
alpinus,  which  also  grows  large  in  the  sea.66 

Occurrence  in  the  Gulf  of  Maine. — When  the 
white  man  first  came  to  New  England  and  to  the 
Maritime  Provinces,  he  found  salmon  in  every 
large  stream  not  barred  by  impassable  falls, 
from  Cape  Sable  to  Cape  Cod;  i.  e.,  in  all  the  Nova 
Scotia  and  New  Brunswick  Rivers,  tributary 
either  to  the  open  Gulf  of  Maine  or  to  the  Bay 
of  Fundy,  and  in  the  following  rivers  in  New 
England:  St.  Croix,  Dennys,  Orange,  East 
Machias,  Pleasant,  Narraguagus,  Union,  Penob- 
scot, St.  George,  Medomak,  Sheepscot,  Andro- 
scoggin, Kennebec,  Royal,  Presumpscot,  Saco, 
Mausam,  Piscataqua,  and  Merrimac.67  One  New 
England  river,  however,  after  another  was  so 
obstructed  by  dams  after  the  beginning  of  the 
past  century,  that  salmon  regularly  entered  only 
the  St.  Croix,  Dennys,  East  Machias,  Machias, 
Penobscot,  Sheepscot,  Kennebec,  and  Andro- 
scoggin by  the  1880's.  The  Kennebec  was  still 
an  important  salmon  river  as  late  as  1895.  But 
by  1925  the  Dennys  and  the  Penobscot  alone, 
of  the  rivers  of  Maine,  saw  regular  runs,  with  a  few 
fish  in  the  St.  Croix  where  pollution  by  sawdust 
was  not  as  bad  then  as  it  had  been,  perhaps  with 
an  occasional  fish  in  other  streams. 

The  fate  of  the  salmon  in  the  Merrimac  M  typi- 
fies its  history  in  the  rivers  from  which  it  is  now 
barred.  Salmon  spawned  plentifully  in  the  upper 
tributaries,  especially  in  the  Pemigewasset,  as  late 
as  1793  (in  1790  the  run  was  so  abundant  in  the 
lower  river  that  60  to  100  a  day  was  the  usual  catch 
with  a  90-yard  seine  near  the  mouth  at  Amesbury) , 
but  the  completion  of  the  dam  at  Lawrence  in  1847 
completely  barred  the  upper  reaches  of  the  river. 
For  some  years  thereafter  salmon  congregated 
below  the  Lawrence  dam  in  spring  and  summer, 
vainly  endeavoring  to  ascend,  but  there  has  beeu 
no  run  of  salmon  in  the  upper  Merrimac  since  1859 
or  1860,  when  the  last  salmon  hatched  above  the 
dam  had  lived  its  span  of  life,  nor  have  any 
spawned  there  since  then  with  the  possible  excep- 
tion of  a  few  that  have  been  lifted  over  the  dam  by 
hand. 

*  Blair  (Res.  Bull.  12,  Dept.  Nat.  Resources  Newfoundland,  1943,  pp.  5-17) 
gives  a  detailed  account  of  the  salmon  rivers  of  the  outer  Labrador  coast, 
Strait  of  Belle  Isle  to  Hamilton  Inlet. 

■  Atkins  (1887,  Fish.  Ind.  U.  S.,  Sect.  5,  vol.  1,  p.  679)  has  collected  much 
Information  on  the  local  history  of  salmon  in  northern  New  England. 

"  Lyman  and  Reed,  Kept.  Comm.  Fish.  Massachusetts  (1865)  1866, 
Senate  Doc.  8,  pp.  36-41. 


126 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


Enough  salmon  to  yield  a  supply  of  eggs  for 
artificial  hatching  continued  to  enter  the  lower 
Merrimac  up  to  1893  and  there  seems  to  have  been 
what  almost  might  be  described  as  a  run  there  in 
1896,  when  salmon  were  seen  leaping  below  the 
Lawrence  dam  nearly  every  day  from  June  10th 
to  July  25th,  often  10  or  20  at  a  time,  and  a  few 
were  lifted  over.  But  we  have  not  learned  of  a 
single  sea-run  salmon  seen  in  the  Merrimac  since 
1901,  though  watch  has  been  kept  for  them  by  the 
wardens  of  the  Massachusetts  Division  of  Fish- 
eries and  Game,8*  and  it  is  not  likely  that  salmon 
would  still  run  in  the  Penobscot  were  it  not  for  the 
artificial  propagation  that  is  carried  on  there  by 
the  U.  S.  Fish  and  Wildlife  Service.  But  the  sal- 
mon situation  now  shows  signs  of  improvement, 
for  the  run  in  the  St.  Croix  has  increased;  salmon 
have  reestablished  themselves  in  the  Narraguagus 
and  provide  sport  there  for  many  anglers  since 
one  obstructing  dam  has  washed  out  and  another 
opened.  Enough  salmon  run  regularly  in  the 
Dennys  to  attract  anglers  and  a  few  also  in  the 
Machias  and  Pleasant  Rivers.  The  Fisheries  Com- 
mission of  the  State  of  Maine,  and  the  U.  S. 
Fish  and  Wildlife  Service,  are  now  concerned  with 
the  possibility  of  improving  the  runs  in  these 
streams,  and  of  reestablishing  runs  of  salmon  in 
other  Maine  rivers. 

Along  the  Canadian  shores  of  our  Gulf  a  few 
salmon  still  run  in  the  Tusket,  Salmon,  and  An- 
napolis Rivers;  many  in  the  Shubenacadie  River 
in  Nova  Scotia,  some  in  the  Petitcodiac,  and 
great  numbers  in  the  St.  John  River  in  New 
Brunswick,  which  still  is  a  famous  salmon  river. 

Movements  in  the  Gulf. — After  the  smolts  reach 
salt  water  they  are  found  for  a  time  in  the  river 
mouths  and  about  estuaries.  No  doubt  the  little 
salmon  (too  small  to  sell)  that  are  caught  in  sum- 
mer and  autumn  in  weirs  at  Matinicus  Island 
have  come  from  the  Penobscot  a  month  or  two 
previous.  They  drop  out  of  sight  in  winter,  as 
do  the  older  and  larger  salmon  as  well.  But 
there  is  no  reason  to  suppose  that  many  of  them 
go  far  out  to  sea  in  the  Gulf.  Odd  salmon 
stray,  it  is  true,  as  much  as  90  to  100  miles  sea- 
ward off  the  outer  coast  of  Nova  Scotia,70  while 


M  A  few  small  "salmon"  reported  of  late  In  the  Merrimac  probably  were 
the  landlocked  form,  running  down  from  tributaries  stocked  with  this  flsh. 

■*  Three  reports  of  salmon  caught  on  Western  Bank  have  appeared  in  the 
daily  press  since  1925  to  our  knowledge,  and  Kendall  (Mem.  Boston  Soc.  Nat. 
Hist.,  vol.  9,  No.  1,  1935,  p.  33)  reports  one  caught  on  La  Have  Bank  100 
miles  from  Halifax,  and  another  60  miles  off  Cape  Sable. 


otter  trawlers  pick  up  odd  salmon  from  time  to 
time  in  the  South  Channel,  and  even  on  Georges 
Bank  up  to  160  miles  or  more  at  sea  from  Cape 
Cod.71  But.  the  great  majority  of  the  salmon 
that  are  caught  in  the  Gulf  are  taken  within  25 
miles  of  the  land. 

The  Gulf  of  Maine  salmon  also  appear  to  con- 
tinue rather  closely  localized  as  a  whole,  not  only 
within  the  coastal  belt,  but  within  the  zone  of 
fresh-water  influence  from  the  particular  rivers  or 
river  systems  from  which  they  come.  So  few,  for 
example,  are  caught  near  Cape  Sable  that  there 
can  be  no  general  movement  around  the  Cape  by 
the  fish  that  spawn  in  the  rivers  of  the  outer  coast 
of  Nova  Scotia.  Most  of  the  fish  that  go  to  sea 
via  Minas  Channel  from  the  Shubenacadie,  and 
the  few  from  smaller  streams  that  discharge  into 
Minas  Basin,  seem  to  remain  along  the  Nova  Scotia 
shore  within  a  distance  of  30  to  40  miles  to  the 
westward.  And  while  tagging  experiments  have 
proved  that  some  of  them  scatter  more  widely;  i.  e. 
to  Cobequid  Bay,  to  the  estuary  of  the  St.  John 
River,  to  the  Annapolis  Basin,  and  to  St.  Mary's 
Bay,  few  of  them  leave  the  Bay  of  Fundy72  (for 
some  that  did,  see  p.  127). 

The  much  more  numerous  salmon  from  the  St. 
John  appear  to  hold  rather  closely  to  the  tongue  of 
low  salinity  that  extends  westerly  from  the  mouth 
of  the  river,  keeping  out  from  the  shore,  for  hardly 
any  salmon  are  caught  either  on  the  New  Bruns- 
wick shore  to  the  eastward,  except  for  a  few  near 
the  head  of  the  Bay  (doubtless  the  product  of  the 
Chignecto  Bay  river  system)  or  farther  west  than 
Point  Lepreau,  or  around  Grand  Manan  Island 
which  stands  directly  in  the  route  of  any  fish  mov- 
ing westward  out  of  the  northern  side  of  the  Bay  of 
Fundy.  Thus  it  appears  that  a  radius  of,  say,  40 
to  50  miles  would  enclose  the  wanderings  of  most  of 
the  St.  John  River  fish. 

The  evident  failure  of  salmon  from  the  St.  John 
to  follow  the  myriads  of  sardine  sized  herring  into 
Passamaquoddy  Bay  is  especially  interesting.  The 
weirs  there  pick  up  a  few  salmon,  the  presence 
of  which  can  be  credited  to  the  small  run  in  the  St. 
Croix  River.  And  the  numbers  of  salmon  that  are 
caught  thence  westward  along  the  coast  of  Maine73 

'■  Kendall  (Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1, 1935.  pp.  31-33)  lists 
a  number  of  such  cases. 

'»  Huntsman,  Ann.  Rept.  Fish.  Ees.  Board  Canada,  (1947)  194S,  p.  37,  and 
unpublished  notes. 

"  The  average  was  only  3,000  pounds  (perhaps  300  flsh)  for  the  years  1939, 
1940,  1943,  1944.    Statistics  are  not  readily  available  for  1941  and  1942. 


FISHES   OF  THE  GULF  OF   MAINE 


127 


are  not  larger  than  can  be  credited  to  such  of  the 
Maine  rivers  as  still  have  runs  of  salmon. 

It  seems  certain,  also,  that  only  odd  salmon  from 
the  Penobscot  and  from  the  rivers  farther  east 
ordinarily  disperse  westward  and  southward  be- 
yond Casco  Bay,  for  while  the  average  catch  for  the 
coast  of  Maine  east  of  that  point  has  averaged 
about  12,000  pounds  (some  1,200  fish)  for  the  10 
most  recent  years  of  record74  combined,  the  cor- 
responding 10-year  average  for  the  whole  western 
side  of  the  Gulf  from  Cape  Elizabeth  to  the  elbow 
of  Cape  Cod  was  only  600  to  700  pounds,  or  some 
60  to  80  fish  at  most,  with  more  than  100  pounds 
reported  in  only  5  of  the  10  years  and  none  in 
3  of  the  years.  Further  evidence  of  a  more  gen- 
eral kind  that  Gulf  of  Maine  salmon  do  not  scatter 
far  as  a  rule  is  that  they  appear  about  the  river 
mouths  in  spring  so  soon  after  the  ice  goes  out  that 
they  cannot  have  come  from  any  great  distance. 

A  few  do  stray  as  far  as  Cape  Cod  Bay  in  most 
years;  witness  catches  of  one  to  5  or  6  fish  (10-55 
pounds)  in  14  out  of  16  years  by  8  traps,  at  North 
Truro,  Cape  Cod,  during  the  period  1935  to  1950, 
in  the  months  of  May,  June,  July,  September,  and 
November.76 

A  year  comes  from  time  to  time  when  a  con- 
siderable number  are  taken  off  the  coast  of 
Massachusetts.  The  most  recently  recorded  in- 
stance of  this  sort  fell  in  1937,  when  floating  traps 
along  the  North  Shore  of  Massachusetts  Bay 
picked  up  4,400  pounds  of  salmon.  All  of  these 
were  taken  close  inshore.  But  the  1,600  or  so 
salmon  (16,050  lb.)  that  were  reported  for  Massa- 
chusetts in  1928  (the  big  year  next  previous) 
seemingly  were  farther  out  at  sea,  for  all  of  them 
either  hooked  on  long  lines  (10,134  lb.),  or  were 
taken  in  otter  trawls.  These  must  have  come  from 
as  far  as  the  Penobscot,  if  not  from  the  Bay  of 
Fundy,  which  is  equally  true  of  the  salmon  that 
are  caught  around  Marthas  Vineyard  from  time 
to  time.78  One,  however,  of  about  10  pounds, 
reported  in  the  North  River,  Marshfield,  in  the 
summer  of  1938,  and  a  few  seen  jumping  in  the 
Parker  River  (also  in  Massachusetts)  in  the  sum- 
mer of  1951,  may  have  been  the  product  of 
attempts  to  stock  these  streams.  Occasional  sal- 
mon that  have  been  taken  along  the  New  Jersey 


N  1933, 1935, 1937, 1938, 1939, 1940, 1943, 1944, 1945,  and  1946. 
"  Information  contributed  by  the  Pond  Village  Cold  Storage  Co. 
"  In  the  spring  of  1915  about  75  (including  fish  up  to  35  lb.)  were  taken  at 
Gay  Head  and  in  the  neighborhood  of  Woods  Hole 


coast  and  off  Delaware77  may  have  been  the 
product  of  attempts  to  stock  the  Hudson. 

Salmon,  also,  of  25  to  50  pounds  that  are 
sometimes  caught  in  Minas  Channel  at  the  head 
of  the  Bay  of  Fundy,  must  come  from  afar,  as 
Dr.  Huntsman  points  out,78  probably  from  the 
Gulf  of  St.  Lawrence,  there  being  no  run  of  fish 
so  heavy  in  any  Bay  of  Fundy  river  or  in  any 
Maine  river. 

It  is  not  astonishing  that  some  salmon  should 
stray  far  afield  in  Gulf  of  Maine  waters,  for 
marked  salmon  have  been  known  to  make  much 
longer  journeys,  elsewhere.  Thus  fish  marked  in 
the  southern  side  of  the  Gulf  of  St.  Lawrence  have 
been  recaught  on  the  north  shore  of  the  Gulf;  in 
Newfoundland;  and  in  the  Strait  of  Belle  Isle.7' 
One  marked  at  Bonavista  on  the  east  coast  of 
Newfoundland  was  retaken  98  days  later  in  the 
Margaree  River,  Cape  Breton  Island,  Nova 
Scotia,  550  miles  away  80  by  the  shortest  possible 
route.  One  marked  in  Minas  Channel  at  the 
head  of  the  Bay  of  Fundy  went  out  around  Nova 
Scotia  to  Chedabucto  Bay  on  the  northeast,  near 
the  Gut  of  Canso,  a  journey  of  at  least  440  miles.81 
Five,  tagged  in  the  Annapolis  River  system,  were 
recaught  on  the  east  coast  of  Newfoundland,  a 
minimum  distance  of  900  miles,  while  a  sixth, 
from  the  same  lot,  was  taken  at  Ramah  on  the 
outer  coast  of  Labrador,  more  than  1,000  miles 
still  farther  away  to  the  northward.82  This  last 
is  the  most  spectacular  case  of  wandering  yet 
reported  for  any  Gulf  of  Maine  or  Gulf  of  St. 
Lawrence  salmon. 

What  is  chiefly  interesting  about  the  large 
catches  that  are  sometimes  made  off  Massachu- 
setts is  their  demonstration  that  so  many  fish  may 
occasionally  wander  so  far  afield.  And  this  ap- 
plies not  only  to  large  salmon  but  to  smolts  in 
their  first  year  at  sea,  for  salmon  so  small  that 
they  must  have  run  down  to  salt  water  but  a  few 
months  previous  have  been  taken  in  Cape  Cod 
Bay  in  October. 

It  is  not  likely  that  these  wandering  salmon 
return  at  all  to  their  home  rivers;  probably  they 

i»  Smith  (Bull.  V.  9.  Fish.  Comm.,  vol.  14,  1896,  p.  99)  reports  salmon 
seined  among  some  mackerel  off  Delaware  in  1893. 

»  Bull.  51.  Biol.  Board  of  Canada,  1936,  p.  9. 

'•  See  Huntsman,  Pub.  Amer.  Assoc.  Adv.  Sci.  8,  1932,  p.  35,  for  summary 
of  these  records. 

"  Huntsman,  Science,  vol.  95,  1947,  p.  381. 

«  Huntsman,  Ann.  Rept.  Fish.  Res.  Bd.  Canada,  (1947)  1948,  p.  37. 

«•  Huntsman,  Science,  vol.  85,  1937,  p.  314;  Pub.  8,  Amer.  Assoc.  Adv. 
Sci.,  1939.  p.  35. 


128 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


are  lost  permanently  from  the  breeding  popula- 
tion. But  the  much  greater  numbers  that  remain 
localized  not  very  far  from  their  parent  streams 
are  believed  to  follow  about  the  same  routes  on 
their  return  journeys  that  they  followed  when 
they  went  to  sea.  Thus,  only  a  few  are  caught 
on  the  Nova  Scotia  shore  between  the  entrance  to 
St.  Mary's  Bay  and  Digby  Gut,  but  fish  en  route 
to  the  Shubenacadie  River  system  are  taken  in 
some  numbers  as  they  follow  the  shore  of  An- 
napolis and  Kings  Counties  (the  Annapolis  River 
also  yields  a  few  salmon  in  its  lower  course,  and 
some  are  taken  in  the  Annapolis  Basin).  Simi- 
larly, salmon  approaching  the  St.  John  River 
strike  the  coast  about  Point  Lepreau  (about  23 
miles  to  the  west)  and  support  an  important 
fishery  from  there  to  the  mouth  of  the  river. 

A  question  closely  bound  to  the  movements  of 
salmon  to  the  sea  is:  what  proportion  of  them 
return  to  spawn  in  the  very  rivers  in  which  they 
were  hatched?  It  seems  demonstrated  by  a 
variety  of  evidence,  especially  by  the  recapture 
of  tagged  fish,  that  the  majority  do  return. 
Huntsman,  for  example,  reports83  an  extraordinary 
instance,  of  a  kelt  taken  from  the  Sackville  River 
on  the  outer  coast  of  Nova  Scotia  that  was  tagged 
and  released  in  the  Shubenacadie  River  system  at 
the  head  of  the  Bay  of  Fundy,  and  then  found  its 
way  out  of  the  Bay,  around  the  Nova  Scotia 
coast,  and  back  again  to  the  Sackville,  where  it 
was  recaptured.  We  can  only  speculate  how  it 
directed  its  course,  and  why  it  did  not  turn  in  to 
the  mouth  of  any  of  the  other  salmon  rivers  it 
passed  en  route.  On  the  other  hand,  marked 
fish  are  sometimes  caught  in  strange  rivers. 
Fish,  for  instance,  that  were  tagged  in  Minas 
Channel  have  been  caught  later  in  the  St.  John 
River.84  And  odd  fish  appear  from  time  to  time 
in  rivers  where  no  salmon  have  been  hatched  for 
many  years  (in  the  Merrimac  for  instance) . 

In  short,  the  parent-stream  theory  does  not 
always  hold.  Probably  the  truth  is  that  while 
most  of  the  fish  never  stray  far  away  and  do 
return  to  the  home  stream,  wanderers  that  chance, 
in  the  spring,  to  be  in  the  physical  state  leading 
to  maturity  may  enter  any  unpolluted  stream 
they  encounter,  no  matter  how  far  from  home. 

Dr.  Huntsman's  studies,  carried  on  through 
many  years,  make  it  increasingly  probable  that 

•»  Ann.  Rept.  Fish.  Res.  Bd.  Canada  (1947)  1948,  p.  33. 

"  Huntsman,  Ann.  Rept.  Fish.  Res.  Bd.  Canada  (1948)  1949,  p.  40. 


the  journeyings  of  our  salmon  in  salt  water  are 
not  the  result  of  purposeful  swimming  in  a  definite 
direction,  but  that  they  tend  to  drift  with  the 
current  as  herring  do  (p.  97),  so  that  the  direction 
in  which  they  travel  depends  chiefly  on  the  depth 
at  which  they  happen  to  be,  in  relation  to  the  dif- 
ferential circulation  of  the  water  at  different 
levels.  If  so,  the  St.  John  River  fish  tend  to  drift 
out  with  the  river  water  as  they  scatter.  And 
most  of  them  do  appear  to  remain  more  or  less 
concentrated  in  the  mid-depths  where  the  princi- 
pal mixing  takes  place  between  the  river  dis- 
charge and  the  water  of  the  open  Bay  of  Fundy, 
some  20  to  30  miles  from  St.  John  Harbor,  living 
where  they  find  an  abundance  of  herring  of  various 
sizes  as  food.  Here  Dr.  Huntsman  85  calculates 
the  space  for  them  is  so  great  that  no  two  of  the 
approximately  50,000  fish  that  comprise  the  total 
yearly  catch  need  be  closer  to  each  other  than 
three-quarters  of  a  mile  in  a  layer  of  water  5  feet 
thick;  so  there  is  no  crowding.  But  the  tagging 
experiments  have  shown  that  the  fish  that  go  to 
sea  from  Minas  Channel,  where  the  outflow  is  not 
so  definitely  localized,  scatter  more  widely,  some  of 
them  drifting  right  around  the  Bay  of  Fundy 
with  the  anti-clockwise  circulation.88 

The  situation  is  not  so  clear  for  the  coast  of 
Maine,  partly  because  of  the  paucity  of  present- 
day  information,  partly  because  the  several 
rivers  there  that  once  had  runs  of  salmon  are  so 
closely  spaced  along  the  coast  that  it  is  not  pos- 
sible to  evaluate  their  individual  contributions  to 
the  yearly  catches. 

With  the  relationship  between  salmon  journeys 
and  water  movements  so  extremely  complex,  all 
we  dare  say  in  this  regard  is  that  the  inshore  drift 
of  the  deeper  layers  (characteristic  of  circulation 
of  the  estuarine  type)  and  the  slackening  of  the 
offshore  drift  of  the  fresher  surface  water  that  is 
to  be  expected  as  the  spring  freshets  diminish, 
may  be  the  cause,  at  least  in  part,  for  bringing 
the  salmon  into  the  estuaries,  and  close  inshore 
elsewhere,  in  spring.  But  the  nature  of  the  stim- 
ulus that  impels  a  salmon  to  enter  fresh  water, 
and  then  fight  his  or  her  way  upstream,  remains 
a  mystery. 

It  is  not  known  whether  all  the  salmon  move 
inshore  in  spring,  or  only  those  that  are  destined 

"  Bulletin  21,  Biol.  Bd.  Canada,  1931,  p.  96. 

M  This  was  shown  by  Huntsman,  Ann.  Rep.  Fish.  Res.  Bd.  Canada  (1947) 
1948,  p.  37. 


FISHES  OF  THE  GULF  OF   MAINE 


129 


to  spawn  that  year,  plus  a  certain  number  of  im- 
mature grilse  that  have  passed  1  year  at  sea.  And 
Dr.  Huntsman87  has  pointed  out  that  the  move- 
ment of  the  salmon  riverward  may  be  very  slow; 
thus  the  salmon  may  take  as  much  as  a  month  to 
cover  the  20-odd  miles  to  the  head  of  tide  in  the 
Petitcodiac  River,  while  some  of  those  that  enter 
the  estuary  of  the  St.  John  River  in  autumn  pass 
the  winter  there  (probably  in  a  lethargic  state) 
before  moving  up  to  the  head  of  tide  80  miles 
distant.  In  any  case,  only  such  fish  as  are  ap- 
proaching sexual  maturity  (irrespective  of  age), 
and  some  immature  female  grilse,  run  far  up  into 
the  rivers;  all  the  others  remain  in  salt  water,  or 
at  most  they  do  not  run  above  the  head  of  tide,  as 
has  often  been  remarked. 

The  majority  of  the  Gulf  of  Maine  salmon  be- 
come "river-mature"  as  it  is  called,  long  before  the 
spawning  season,  for  while  none  of  them  spawn 
before  October,  some  of  them  enter  fresh  water  as 
early  as  March  and  April.  But  the  chief  runs  come 
later,  varying  in  date,  not  only  from  river  to  river, 
but  from  year  to  year  in  a  given  river.  In  the 
Penobscot,  some  fish  may  enter  in  March ;  they  are 
to  be  expected  in  the  lower  reaches  after  the  first 
week  in  April;  more  come  in  May,  perhaps  two- 
thirds  in  June,  with  a  few  fish  entering  later  still. 
Available  information  is  to  the  effect  that  few  enter 
the  Narraguagus  and  Dennys  Rivers  until  well 
into  May,  the  chief  runs  there  coming  in  June,  with 
some  entering  as  late  as  September.  We  have  not 
been  able  to  obtain  definite  dates  for  the  spring 
and  early  summer  runs  in  the  St.  John  River.  But 
it  seems  that  salmon  continue  to  enter  the  latter 
until  well  into  the  autumn,  judging  from  catches 
of  fish  so  fat  that  they  must  have  come  in  recently 
from  the  sea.  Salmon  enter  other  streams  tribu- 
tary to  the  Bay  of  Fundy  from  May  on.  As  a 
rule  the  large  salmon  come  earliest,  the  grilse  not 
until  later,  probably  because  it  is  not  until  later 
that  the  latter  have  reached  the  degree  of  fatness 
associated  with  river  maturity.  Accordingly,  the 
heaviest  runs  in  the  Shubenacadie,  mostly  grilse 
(p.  130.),  are  said  to  come  from  August  until  late 
in  the  autumn. 

Every  salmon  fisherman  is  familiar  with  the  fact 
that  salmon  enter  in  "runs"  that  are  spaced  irreg- 
ularly in  time,  and  that  vary  in  date  from  year 
to  year,  depending  on  the  height  of  water  in  the 

"  Progress  Report,  Atlantic  stations,  Biol.  Bd.  Canada.  8,  1933,  p.  6;  and 
unpublished  notes. 

210941—53 10 


river  and  on  the  strength  of  the  current.  Freshets 
tend  to  bring  them  in;  if  the  current  becomes  too 
strong  they  simply  hold  position,  to  breast  it 
again  as  the  flow  slackens.  The  fish  that  are  in 
the  estuary  remain  there  during  the  periods  be- 
tween freshets,  waiting,  as  it  were,  for  the  message 
from  upstream  that  starts  them  on  their  way. 
And  the  salmon  within  the  river  are  similarly 
quiescent  during  periods  of  low  water  and  weak 
current.  This  is  the  chief  reason  why  salmon 
angling  is  so  uncertain  a  sport,  even  in  the  best  of 
rivers. 

A  good  deal  of  discussion  has  centered  about  the 
question  whether  the  earliest  fish  stay  in  fresh 
water  from  then  until  spawning  time  (a  matter 
of  6  months)  or  whether  there  is  more  or  less  move- 
ment in  and  out  of  the  river  mouths  at  the  begin- 
ning of  the  season.  The  latter  view  may  be  cor- 
rect for  the  small  streams,  but  it  seems  safe  to 
say  that  after  the  run  is  well  under  way  in  late 
May  or  early  June  no  fish  return  from  fresh  to  salt 
water  until  autumn.  Tagging  experiments  carried 
out  in  Canadian  rivers  have  also  yielded  the  very 
interesting  information  that  no  matter  when  a 
salmon  runs  upstream  in  one  year,  it  may  do  so 
either  early  or  late  in  the  next.88 

It  is  a  matter  of  common  knowledge  that  salmon 
average  larger  in  some  rivers  than  in  others,  and 
growth  studies  based  on  the  scales  have  shown  that 
these  differences  are  due  chiefly  to  the  average 
ages  of  the  fish  that  enter.  In  the  St.  John,  as 
Huntsman  has  pointed  out,89  there  are  three  prin- 
cipal groups  of  salmon:  (a)  male  grilse,  averaging 
about  6  pounds,  that  are  mature  and  fated  to 
breed  that  same  autumn;  (b)  the  ordinary  spawn- 
ers  that  have  passed  two  years  or  more  at  sea; 
these  average  10  to  15  pounds  in  weight  and  enter 
from  May  to  August,  the  late  comers  running 
heavier  than  the  early  comers;  most  of  them  are 
virgins,  but  some  of  them  have  already  spawned 
once  or  twice;  (c)  immature  female  grilse,  averag- 
ing about  9  pounds,  that  enter  from  November  to 
January.  Few,  however,  return  to  spawn  in  the 
rivers  of  Maine  until  they  have  passed  2  years  at 
sea ;  not  more  than  3  or  4  grilse  to  70  adults  were 
taken  in  the  St.  Croix,  for  example,  when  there 
still  was  a  good  run  there,  and  not  more  than  1 


*  Fifty-fifth  Annual  Report  of  the  Fisheries  Branch,  Department  of  Marine 
and  Fisheries,  Canada,  (1921-22)  1922,  p.  19. 
■  Nature,  vol.  141, 1938,  p.  421;  Pub.  8,  Amer.  Assoc.  Adv.  Sci ,  1939,  p.  34. 


130 


FISHERY  BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


grilse  to  500  adults  in  the  Dennys  and  Penobscot 
Rivers.90 

The  average  weight  of  the  salmon  caught  in 
the  Penobscot  was  about  UK  pounds  in  1905 
(6,378  fish),  9  to  10  pounds  in  1919  and  1920 
(3,920  fish),91  or  a  little  less  than  in  the  St.  John. 
The  heaviest  Penobscot  fish  of  which  we  found 
definite  record  of  late  years  weighed  a  little  more 
than  35  pounds.92  The  fish  in  the  rivers  flowing 
into  the  head  of  the  Bay  of  Fundy  run  much 
smaller,  as  Perley  pointed  out  long  ago,  and  recent 
studies  show  that  most  of  them  spawn  first  as 
grilse,  i.  e.,  after  only  one  year  at  sea;  a  few, 
having  spawned  after  one  year  at  sea,  return  to 
spawn  again  a  year  later;  and  the  percentage  of 
larger  and  older  fish  is  very  small  there.  This, 
Huntsman  points  out,93  contrasts  with  the  preva- 
lent 6-year-old  fish  m  the  Miramichi,  which  dis- 
charges into  the  southern  side  of  the  Gulf  of 
St.  Lawrence,  and  with  7-  or  8-year-old  fish  in 
the  Grand  Cascapedia,  tributary  to  the  Bay  of 
Chaleur.  Various  explanations  have  been  ad- 
vanced to  account  for  these  differences  from  river 
to  river,  none  of  them  convincing  in  our  opinion. 

It  also  appears  to  be  true  (as  often  stated) 
that  a  larger  proportion  of  the  salmon  are  annual 
spawners  in  small  streams,  where  most  of  the 
spent  fish  drop  downstream  again  soon  after 
spawning,  than  in  large  rivers  where  many  of 
these  "kelts"  remain  in  fresh  water  over  the 
winter.  A  plausible  explanation  is  that  kelts 
that  return  to  the  sea  immediately  after  spawning 
have  less  opportunity  to  grow  (though  they  recover 
condition  sufficiently  to  spawn  again  the  following 
summer)  than  such  as  await  the  spring  to  go 
downstream,  and  that  spend  a  whole  year  at 
sea  instead  of  one  winter  only  between  two  suc- 
cesive  spawnings.  This,  however,  does  not  ac- 
count for  the  fact  that  it  is  almost  invariably 
the  large  rivers  that  yield  the  very  large  maiden 
fish  that  have  spent  4  years  at  sea,  or  more. 

Abundance. — The  early  extirpation  of  salmon 
from  the*Merrimac,  Saco,  Kennebec  system,  and 
various  rivers  to  the  eastward  naturally  resulted 
in  a  great  decrease  in  the  abundance  of  salmon 

»  See  Kendall  (Mem.  Boston  Soe.  Nat.  Hist.,  vol.  9,  No.  1, 1935,  pp.  58-60) 
for  age  determinations  of  Penobscot  salmon. 
•'  Radclifle.  Rept.  U.  S.  Comm.  Fish.  (1921)  1922,  p.  146. 
*  Kendall.  Mem.  Boston  Soc.  Nat.  Hist.,  vol.  9,  No.  1,  1935,  p.  32. 
■  Bull.  21.  Biol.  Board  Canada.  1931.  p.  19. 


in  the  open  Gulf,  clearly  reflected  in  the  catches. 
Data  are  not  available  for  early  years  when  all 
the  rivers  still  offered  free  access.  But  the  yearly 
catch  had  been  reduced  to  about  100  to  500  fish 
in  the  St.  Croix  by  about  1887;  200  to  1,000 
each  in  the  Dennys  and  in  the  Kennebec,  and 
5,000  to  15,000  in  the  Penobscot.  The  catch 
along  the  Maine  coast,  which  had  been  a  little 
more  than  150,000  pounds  in  1889  (more  than 
seven-eighths  of  this  in  or  about  the  approaches 
to  the  Penobscot),  was  only  about  86,000  pounds 
in  1905  (of  this  74,000  lb.,  or  6,378  fish  from  the 
Penobscot);  was  about  20,000  pounds  in  1919 
(13,557  lb.  or  1,322  fish  from  the  Penobscot); 
and  was  only  14,744  pounds  (12,700  lb.  or  1,221 
fish  from  the  Penobscot)  in  1928.  As  70  to  90 
percent  of  the  Maine  catch  comes  from  Penobscot 
River  or  Bay,  the  following  table  of  salmon 
caught  there  in  certain  years  from  1896  to  1928 
is  pertinent: 


Year 

Number 

offish 

Pounds 

Year 

Number 
offish 

Pound! 

1896. . 

..  6,404 

80,  225 

1918. 

—   1,653 

17,  212 

1898. . 

..  3,225 

42,  560 

1919. 

...   1,322 

13,  557 

1901- . 

..  6,821 

86,  055 

1920. 

— .   1,598 

15,  135 

1903.. 

..  4,859 

67,  470 

1928. 

...   1,221 

12,  700 

1905. . 

..  6,378 

74,  158 

The  Maine  catch  then  increased  again  to  about 
88,000  pounds  in  1930  and  to  about  70,000  in  1931, 
suggesting  a  better  run  in  the  Penobscot,  and  var- 
ied between  16,000  and  40,000  pounds  through  the 
period  1932-1938.94  But  the  average  reported 
catch  for  Maine  for  the  period  1939  to  1947 95  was 
only  about  3,600  pounds  (maximum  9,300,  min- 
imum 600),  the  average  Massachusetts  catch  for 
the  same  period  only  about  100  pounds  (maxi- 
mum 400,  minimum  0).  Thus  the  output  of 
salmon  from  the  rivers  of  Maine  (none  from  the 
rivers  of  Massachusetts)  has  been  only  about  one- 
fiftieth  as  great  during  the  past  few  years  as  it 
was  some  60  years  ago. 

The  numbers  of  salmon  have  held  up  much 
better  in  the  Canadian  waters  of  the  Gulf,  thanks 
to  wise  measures  of  conservation  such  as  limiting 
netting  at  the  mouths  of  the  rivers,  and  keeping 
the  streams  free  for  access  by  fishways  at  the  dams. 
The  average  yearly  catches,  from  1870  to  1946, 


«  No  data  are  available  for  1934  or  1936. 
-  No  data  for  1941. 


FISHES   OF  THE  GULF   OF   MAINE 


131 


were  as  follows  for  the  west  coast  of  Nova  Scotia 
and  for  the  Bay  of  Fundy  combined: 


Years  Pounds 

1870-1879 655,  200 

1880-1880 292,700 

1890-1899 634,000 

1900-1909. 576,800 


Years  Pounds 

1910-1919 540,000 

1920-1929 470,300 

1930-1939.. 424,000 

1940-1946 278,000 


The  Canadian  catch  in  the  open  Gulf  and  in  the 
Bay  of  Fundy  may  be  expected  to  run  about 
400,000  to  600,000  pounds  at  the  present  time, 
taking  one  year  with  another,  or  something  like 
40,000  to  60,000  fish,  which  is  perhaps  100  times 
as  great  as  that  for  the  entire  coastline  of  Maine 
and  of  Massachusetts.  And  the  distribution  of  the 
catches  shows  that  the  St.  John  River  contributes 
something  like  four-fifths  of  this,  or  a  yearly  aver- 
age of  some  50,000  fish,88  contrasting  with  only  a 
few  hundred  fish  for  the  Penobscot  in  a  poor  year, 
and  perhaps  up  to  8,000  in  a  good. 

Salmon  anglers  are  only  too  familiar  with  the 
fact  that  the  number  of  fish  that  enter  even  the 
best  of  salmon  rivers  is  much  smaller  in  some 
years  than  in  others.  During  the  16-year  period, 
1931-1946,  the  commercial  catches  reported  for 
St.  John  Harbor  and  St.  John  River  (best  salmon 
river  tributary  to  the  Gulf  of  Maine)  were  good 
in  1931  (164,000  lbs.);  in  1935  (149,300  lbs.); 
in  1936  (148,600  lbs.);  in  1937  (172,700  lbs.);  and 
in  1943  (157,500  lbs.);  but  were  poor  in  1939 
(48,500  lbs.);  in  1945  (60,000  lbs.)  and  in  1946 
(54,500  lbs.).  The  yearly  average  for  this  period 
was  116,000  pounds. 

'•  Huntsman  (Bull.  21,  Biol.  Board  Canada,  1931)  has  made  a  very  inter- 
esting analysis  of  catches  for  the  Bay  of  Fundy  as  a  whole,  as  well  as  for  the 
St.  John,  for  the  Chignecto  system,  and  for  the  Minas  system,  separately. 


In  the  Minas  system  the  fishery  produced  as 
much  as  383,800  pounds  in  1907,  283,400  pounds 
in  1917,  and  226,500  pounds  in  1918;  but  since 
then,  up  to  1946,  the  best  catches  have  been  only 
160,700  pounds  in  1919,  165,100  pounds  in  1923, 
and  143,300  pounds  in  1925,  while  the  poorest 
were  28,100  pounds  in  1938  and  26,600  pounds  in 
1945.  The  average  yearly  catch  from  1917  to 
1930  was  133,000  pounds,  and  from  1931  to  1946, 
48,000  pounds. 

The  reader  will  notice  at  once  that  the  big  years 
have  not  been  the  same  for  these  two  bodies  of 
salmon.  It  seems  sufficiently  established  that 
yearly  and  regional  differences,  such  as  these, 
result  in  the  main  from  corresponding  differences 
in  the  numbers  of  smolts  that  reach  salt  water  in 
any  given  year.  And  recent  investigations  in 
Canadian  waters  make  it  likely  that  the  factor 
chiefly  responsible  is  the  height  of  the  water  from 
summer  to  summer,  or  over  periods  of  several 
summers,  which  of  course  reflects  the  yearly  vari- 
ations in  rainfall.  If  the  water  is  high  the  pan- 
are  protected  from  the  birds  that  prey  upon  them 
and  are  more  easily  able  to  escape  the  trout,  so 
that  many  survive  to  descend  to  the  sea  and  to 
return  one,  two,  or  three  years  later.  If  the  water 
in  the  river  is  low  the  parr  are  more  at  the  mercy 
of  kingfishers,  megansers,  and  trout,  so  that  fewer 
of  them  live  to  reach  salt  water,  and  there  are 
fewer  of  them  to  return  as  grilse  or  as  older  fish. 

Humpback     salmon     Oncorhynchus     gorbuscha 
(Walbaum)  1792 

Jordan  and  Evermann,  1896-1900,  p.  478. 


Figure  55. — Humpback  salmon  {Oncorhynchus  gorbuscha). 


132 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE    SERVICE 


Description. — The  humpback  is  of  the  familiar 
salmon  outline  while  living  in  the  sea,  the  body 
being  deeper  than  thick,  with  rounded  belly.  The 
head  is  naked  but  the  body  is  covered  with  scales 
large  enough  to  be  seen  easily.  The  dorsal  fin 
stands  about  midway  of  the  body  above  the  ven- 
trals,  and  the  flaplike  adipose  fin  is  over  the  rear 
end  of  the  anal  fin.  It  agrees  so  closely  with  the 
Atlantic  salmon  in  all  this  that  the  one  might 
easily  be  taken  for  the  other,  were  it  not  that  the 
anal  fin  of  the  humpback  invariably  has  14  rays  or 
more,  whereas  that  of  the  Atlantic  salmon  has 
only  about  9  rays.  Also,  the  humpback  is  a 
stouter-bodied  fish  than  the  Atlantic  salmon. 
The  male  humpback  (like  all  the  Pacific  salmons, 
and  the  Atlantic  salmon  to  a  lesser  degree)  under- 
goes a  very  noticeable  change  in  form  in  the 
spawning  season,  with  the  body  deepening  and 
developing  a  prominent  hump  in  front  of  the  dor- 
sal fin;  the  jaws  elongating  and  becoming  hooked 
at  the  tip  and  the  teeth  increasing  in  size. 

Color. — The  back  and  tail  of  the  humpback  are 
bottle  green  with  poorly  defined  black  spots,  while 
it  is  in  the  sea.  These  spots  are  particularly  con- 
spicuous on  the  tail,  where  they  are  oval  in  outline 
and  as  much  as  a  third  of  an  inch  in  longest  diam- 
eter. These  spots  are  one  of  the  distinctive  marks, 
whereby  the  humpback  can  be  distinguished  from 
all  other  salmons.  The  sides  and  belly  are  sil- 
very, with  a  faint  pinkish  tinge.  Young  hump- 
backs are  unique  among  salmon  in  being  of  prac- 
tically adult  coloration  without  "parr"  marks 
(p.  122). 

Size. — The  humpback  is  the  smallest  of  the 
Pacific  salmons  and  much  smaller  than  the  Atlan- 
tic salmon,  adults  averaging  only  about  5% 
pounds  in  weight  and  20  to  25  inches  in  length. 
Males  weigh  to  about  11  pounds  and  females  to 
about  7%  pounds. 

General  range.- — Pacific  coast  of  North  America 
and  of  northern  Asia,  from  Oregon  northward  on 
the  American  side.  This  is  the  most  abundant 
salmon  in  Alaska.  It  runs  up  fresh  rivers  to 
spawn,  which  it  does  but  once  and  then  dies. 
It  has  been  introduced  in  the  rivers  of  Maine. 

Occurrence  in  the  Gulf  of  Maine. — The  history 
of  the  introduction  of  this  west  coast  salmon  to 
New  England  waters  is  as  follows: 

Humpback  salmon  eggs  seem  first  to  have  been 
planted  in  Maine  rivers  in  1906.  In  the  autumn 
of  1913  a  large  consignment  of  humpback  eggs 


was  shipped  to  the  Craig  Brook  and  Green  Lake 
(Maine)  hatcheries,  and  the  approximately  7,000,- 
000  fingerlings  hatched  therefrom  were  distributed 
in  the  Penobscot,  Androscoggin,  Damariscotta, 
Dennys,  Pleasant,  Union,  Medomak,  Georges, 
and  St.  Croix  Rivers.  A  year  later  some  5,000,000 
young  fish  were  liberated.  A  third  plant  was 
made  in  1915;  a  fourth  of  6,235,808  fingerlings  in 
1916;  and  a  fifth  of  about  1,000,000  in  the  Dennys 
and  Pembroke  Rivers  in  1917.97 

The  results  of  this  attempt  at  acclimatization 
were  first  seen  in  the  summer  and  fall  of  1915 
when  fishermen  reported  large  numbers  of  mature 
humpbacks  along  the  Maine  coast,  and  when 
humpbacks  ran  in  the  Dennys  River  (where  many 
were  caught)  from  August  15  until  September  24, 
some  probably  spawning  there,  for  the  bodies  of 
spent  fish  were  seen  drifting  downstream.  Hump- 
backs again  entered  the  Pembroke  and  Dennys 
Rivers  during  August,  September,  and  October  of 
1917  with  a  few  reported  from  the  Penobscot,  St. 
Georges,  Medomak,  and  St.  Croix,  the  result  of 
the  plant  of  1915.  And  at  least  2,000  mature  fish 
were  seen  that  summer  in  the  Dennys  alone,  where 
many  were  caught  averaging  about  5  pounds,  and 
one  as  heavy  as  10  pounds  9  ounces.  Definite  in- 
formation is  lacking  for  1918.  But  even  larger 
numbers  entered  the  Dennys  and  Pembroke  Rivers 
in  the  autumn  of  1919  than  in  1917,  with  smaller 
runs  in  the  Penobscot,  Machias,  St.  Croix,  and 
Medomak  Rivers.  Enough  spawned  that  year  in 
the  Dennys  and  Pembroke  Rivers  for  the  fish- 
culturists  of  the  Bureau  of  Fisheries  to  artificially 
fertilize  half  a  million  eggs  there.  And  hump- 
backs were  caught  in  the  weirs  in  Passamoquoddy 
and  Cobscook  Bays  during  that  season. 

Adult  fish  were  taken  again  in  the  weirs  in 
1920,98  and  one  fish  was  caught  in  a  weir  as  far 
from  its  native  river  as  Lanesville,  Mass.  (near 
Cape  Ann)  "  at  some  time  during  the  summer  of 
1921. 

Large  numbers  of  eggs  were  collected  again  from 
wild  fish  between  1922  and  1926,  the  resultant  fry 
being  returned  to  the  Dennys  and  other  rivers 
nearby.  Artificial  propagation  was  abandoned 
then,  for  it  seemed  that  the  species  was  estab- 

«  More  detailed  accounts  of  these  and  successive  plantings  will  be  found 
in  tho  annual  reports  of  the  U.  S.  Commissioner  of  Fisheries  for  the  years 
1914  to  1928. 

•>  Reported  catch,  Washington  County,  Maine,  1920,  310  pounds. 

« It  was  forwarded  to  the  Massachusetts  Commissioners  as  reported  by 
C.  E.  Orant  of  Gloucester. 


FISHES   OF  THE  GULF   OF   MAINE 


133 


lished.1  But  natural  reproduction  seems  not  to 
have  been  successful  enough  for  the  humpback  to 
maintain  itself  in  the  few  Maine  rivers  open  to  it, 
much  less  to  increase  in  numbers,  for  very  few  have 
been  reported  since  about  1926  or  1927,  and  none 
that  we  have  heard  of  for  some  years  past. 

Silver  salmon  Oncorhynchus  kisutch 
(Walbaum)  1792 

COHO    SALMON 
Jordan  and  Evermann,  1896-1900,  p.  480. 

Description. — The  silver  salmon  resembles  a 
rather  stout  Atlantic  salmon  closely  in  its  general 
shape,  also  in  the  relative  size  and  position  of  its 
fins,  and  in  their  shapes.  But  a  safe  morphologi- 
cal criterion  for  distinguishing  the  one  from  the 
other  is  that  the  silver  always  has  at  least  12 
rays  in  its  anal  fin,  and  some  of  them  have  as 
many  as  17,  whereas  most  of  the  Atlantic  salmons 
have  only  8  or  9  anal  rays,  and  never  more  than 
10.  The  color  is  a  help  also,  in  this  connection, 
for  while  a  silver  is  silvery  down  its  sides,  like  an 
Atlantic  salmon,  it  is  more  closely  sprinkled  with 
small  black  spots  along  its  back  and  on  the  upper 
part  of  its  tail  fin  than  is  an  Atlantic  salmon. 
These  spots,  too,  are  always  roundish  or  oval  in 
a  silver,  never  in  the  form  of  crosses.  On  the 
other  hand,  the  black  spots  are  much  smaller  and 
much  less  conspicuous  on  a  silver  salmon  than  on 
a  humpback,  and  the  lower  half  of  the  tail  fin, 
which  is  as  conspicuously  spotted  as  the  upper 
half  on  a  humpback,  usually  has  no  spots  on  a 
silver  salmon. 

Size. — Up  to  3  feet  in  length. 

General  range,  habits,  and  occurrence  in  the  Gulf 
of  Maine. — The  native  range  of  the  silver  salmon 
is  from  northern  California  to  northwestern  Alaska, 
where  it  is  an  important  food  fish,  and  where 
anglers  take  many  of  them,  both  by  trolling  and  by 
fly  fishing.     Like  other  Pacific  salmons,  it  runs 

i  Rept.  of  U.  8.  Comm.  Fish.  (1928),  Pt.  1, 1929,  p.  379. 


up  into  fresh  streams  to  spawn,  dying  thereafter- 
Most  of  the  young  remain  about  one  year  in 
fresh  water,  but  a  few  do  not  move  out  to  sea 
until  they  are  in  their  third  year.  Most  of  them 
return  to  fresh  water  at  the  end  of  the  third 
summer  at  sea,  a  few,  however,  by  the  end  of  the 
second  summer  in  salt  water,  a  few  others  not 
until  the  fourth  summer. 

Our  only  reason  for  mentioning  the  silver  salmon 
is  that  a  plant  of  its  fry  and  fingerlings  that  was 
made  in  the  Duck  Trap  stream,  tributary  to  the 
western  side  of  Penobscot  Bay,  near  Lincolnville, 
Maine,  resulted  in  the  return  of  150  mature  fish 
to  Duck  Trap  stream  in  1944,  and  perhaps  of 
more  of  them.  But  nothing  more  was  heard  of 
them  thereafter,  and  no  returns  have  been  re- 
ported up  to  this  writing  (Nov.  1,  1951)  from 
other  plants  that  were  made  in  Maine  waters  2 
in  1948. 

THE  SMELTS.     FAMILY  OSMERIDAE 

The  smelts  are  small  salmons  in  all  essential 
respects,  except  that  their  stomach  has  few  pyloric 
caecae,  or  none,  whereas  there  are  large  numbers 
of  such  caecae  in  their  larger  relatives  of  the  salmon 
family.  However,  it  is  not  necessary  to  look  so 
deeply  to  learn  whether  a  fish  be  smelt  or  very 
young  salmon,  for  the  former  all  have  pointed 
noses  and  are  very  slender,  whereas  the  young  of 
our  four  salt-water  salmons — humpback,  silver 
Atlantic,  and  sea  trout — are  much  stouter,  with 
blunt  noses.  In  most  cases,  too,  the  shape  of 
the  tail  would  suffice  of  itself  to  separate  smelt 
from  salmon  smolt,  for  it  is  never  as  deeply  forked 
in  the  latter  as  in  the  smelts. 

Two  smelt  fishes  occur  in  the  Gulf  of  Maine: 
the  smelt  (very  common),  and  the  capelin  (a 
sporadic  visitor  from  the  north).  The  argentine 
(p.  139)  is  so  closely  related  to  the  smelts  that  it  is 
included  in  the  following  key. 

» In  Tunk  stream,  Duck  Trap  stream,  Chandler  River,  and  Bald  Hill 
Cove  Brook. 


KEY  TO  GULF  OF  MAINE  SMELTS  AND  ARGENTINES 

The  dorsal  fin  originates  over  the  tips  of  the  pectorals;  the  mouth  is  very  small Argentine  p.  139 

The  dorsal  fin  situated  far  behind  the  pectorals;  the  mouth  is  large 2 

Upper  jaw  almost  as  long  as  lower;  teeth  large;  there  is  a  group  of  strong  fangs  on  the  tongue;  the  pectoral  fins  have  12 

rays  or  fewer.. Smelt,  p.  135 

Lower  jaw  much  longer  than  upper;  teeth  so  small  as  hardly  to  be  visible;  no  fangs  on  tongue;  the  pectoral  fins  have 

15  to  20  rays.. Capelin,  p.   134 


134 


FISHERY  BULLETIN   OF  THE    FISH  AND  WILDLIFE   SERVICE 


Capelin  Mallotus  villosus  (Miiller)  1777 
Jordan  and  Evermann,  1896-1900,  p.  520. 

Description. — The  capelin  is  an  even  slenderer 
fish  than  the  smelt,  its  body  being  only  about  one- 
sixth  to  one-seventh  as  deep  and  about  one- 
twelfth  as  thick  as  it  is  long,  and  of  nearly  uniform 
depth  from  gill  cover  to  anal  fin  (except  in  the  case 
of  females  when  their  abdomens  are  distended 
with  spawn),  whereas  the  smelt  is  usually  deepest 
about  its  mid -length  (at  least  if  the  fish  is  fat), 
which  gives  the  two  species  characteristically 
different  aspects.  The  head  of  the  capelin  is 
pointed  like  that  of  the  smelt,  the  mouth  gaping 
back  to  below  the  center  of  the  very  large  eye  with 
the  tip  of  the  lower  jaw  projecting  noticeably  be- 
yond the  upper.  The  scales  are  minute,  much 
smaller  than  those  of  the  smelt  and  more  numerous 
(about  200  per  row  on  the  sides  of  the  body) ;  the 
teeth  so  small  as  to  be  hardly  visible  to  the  naked 
eye,  and  the  tongue  fangs,  so  characteristic  of  the 
smelt  (p.  135),  are  lacking  here.  The  outline  of  the 
adipose  fin  likewise  helps  separate  capelin  from 
smelt,  for  it  is  low  in  the  former  and  about  half 
as  long  as  the  anal,  but  short  and  high  in  the 
latter.  The  pectoral  of  the  capelin  is  broader  also, 
usually  with  15  or  more  rays. 

The  capelin  exhibits  a  pronounced  sexual 
dimorphism ;  the  male  has  much  the  longer  pectoral 
fins;  and  the  base  of  his  anal  is  elevated  on  a  pro- 
nounced hump,  whereas  it  follows  the  general  out- 
fine  of  the  belly  in  the  female.  In  males,  too,  the 
scales  in  one  of  the  longitudinal  rows  immediately 
above  the  lateral  fine,  and  in  another  row  along 
each  side  of  the  belly,  are  pointed,  distinctly  larger 
than  the  other  scales,  and  become  longer  still  at 
spawning  time  when  each  pushes  up  the  skin  as  a 


finger-like  process;  they  form  four  ridges  that  are 
very  evident  when  the  fish  is  held  in  the  hand. 

Color. — The  capelin  is  transparent  olive  to  bottle 
green  above,  like  a  smelt,  but  its  sides  are  uni- 
formly silvery  below  the  lateral  line  and  the  scales 
are  dotted  at  the  margins  with  minute  dusky 
specks  (in  the  smelt  there  is  a  distinct  silvery  band 
on  each  side) ;  the  belly  is  white.  Back  and  head 
darken  at  spawning  time. 

Size. — Few  capelin  are  more  than  6%  to  7% 
inches  long. 

Habits.3 — Capelin  are  most  in  evidence  during 
the  spawning  season,  when  they  come  inshore  in 
multitudes  along  arctic-subarctic  coasts.  They 
spawn  on  gravel  or  pebbly  bottom,  chiefly  close 
below  tideline,  many  of  them  in  the  wash  of  the 
waves  in  the  beach;  many  are  stranded  then 
on  the  beach  between  waves.  But  eggs  have  also 
been  reported  from  as  deep  as  35  to  40  fathoms. 
Each  female  while  spawning  is  accompanied 
by  two  males  that  crowd  her  between  them; 
but  she  may  have  only  one  companion.4  Spawn- 
ing takes  place  chiefly  at  temperatures  of  43°  to 
50°  F.  (6°-10°  C.)  and  more  actively  by  night 
than  by  day. 

The  eggs  are  reddish,  about  1/25-inch  (1  mm.) 
in  diameter,  and  so  sticky  that  they  cling  to  each 
other  like  herring  eggs,  and  to  the  gravel  and 
pebbles  with  which  they  are  intermingled  by  the 
swash  of  the  waves.  They  hatch  in  about  15 
days  at  a  temperature  of  50°  F.  (10°  C).  And 
they  will  tolerate  a  salinity  as  low  as  7  per  mille, 

•  Interesting  accounts  of  the  habits  of  the  capelin  and  of  Its  rate  of  growth 
In  Newfoundland  waters  have  been  given  recently  by  Jeffers  (Ann.  Rept. 
Biol.  Board  Canada  (1930),  1931,  pp.  7-18);  by  Sleggs  (Rept.  Newfoundland 
Fish.  Res.  Comm.,  1,  No.  3, 1933);  and  by  Templeman  (Bull.  Newfoundland 
Government  Lab.,  17  (Research),  1948. 

•  According  to  Lanman,  Rept.  V.  S.  Comm.  Fish.  (1872-1873)  1874,  p.  225. 


Figure  56.— Capelin  (Mallotus  villosus),  Grosswater  Bay  specimen.     From  Goode.     Drawing  by  H.  L.  Todd. 


FISHES  OF  THE  GULF  OF   MAINE 


135 


as  Dr.  Jeff  era  writes  us.  The  larvae,  described 
as  5  to  7  mm.  long  at  hatching,  are  very  slender 
and  resemble  those  of  smelt,  herring,  and  launce 
so  closely  that  identification  is  a  matter  for  the 
expert.  In  any  case,  capelin  are  encountered  so 
seldom  in  our  Gulf  that  their  larvae  are  not  apt 
to  be  seen  there.8 

Along  the  coasts  of  Newfoundland,  capelin 
spawn  chiefly  in  June  and  July,  and  we  have  found 
them  doing  so  in  multitudes  along  the  outer 
Labrador  coast  in  July.  Probably  any  spawning 
that  may  take  place  within  the  limits  of  our  Gulf 
would  fall  in  May  at  latest,  to  judge  from  water 
temperatures. 

The  capelin  so  seldom  appears  in  the  Gulf  of 
Maine  that  we  need  only  add  that  it  is  a  fish  of  the 
high  seas  frequently  encountered  far  out  from  land, 
coming  inshore  only  to  spawn  and  then  as  a  rule 
moving  out  again;  that  it  travels  in  vast  schools 
at  spawning  time  (when  it  often  strands  on  the 
beach  in  countless  multitudes).  It  is  the  chief  bait 
fish  of  Arctic  seas,  preyed  upon  by  whales  and  by 
every  predaceous  fish,  particularly  by  cod, 
which  are  often  seen  pursuing  the  capelin  at  the 
surface  in  northern  waters.  Capelin  themselves 
feed  chiefly  on  small  crustaceans,  particularly 
on  copepods,  on  euphausiid  shrimps,  and  on  am- 
phipods.  It  is  also  known  to  devour  its  own 
eggs.  We  can  bear  witness  that  the  capelin  is 
a  delicious  little  fish  on  the  table. 

General  range. — Boreal-Arctic  seas,  south  to  the 
coast  of  Maine8  on  the  Atlantic  coast  of  America. 

Occurence  in  the  Gulf  of  Maine. — The  capelin 
is  a  sub-Arctic  fieh  that  visits  the  Gulf  of  Maine 
occasionally;  chiefly  the  eastern  side  as  might  be 
expected  since  it  comes  from  the  north. 

Dr.  Hunstman  writes: 7 

In  the  Gulf  of  St.  Lawrence  it  occurs  abundantly  in 
limited  areas,  which  shift  somewhat  from  year  to  year. 
It  occurs  periodically  in  similar  limited  areas  farther  south. 
The  southeastern  corner  of  Cape  Breton  is  the  center  of 
such  an  area,  where  large  quantities  were  taken  in  1917. 
Halifax  is  the  center  of  another  area,  where,  however,  it 
is  more  rare.  In  1916  it  was  abundant  at  Sambro,  near 
Halifax.  The  next  area  is  in  the  Bay  of  Fundy,  where 
they  have,  exceptionally,  been  taken  in  large  quantities 
at  long  intervals. 


•  Tompleman  (Bull.  Newfoundland  Government  Lab.,  17,  Res.,  1948> 
figs.  18-20)  gives  a  series  of  eicellent  illustrations  of  capelin  larvae  at  different 
stages  of  growth. 

8  According  to  Jordan  and  Evermarm  the  capelin  finds  its  southern  limit 
at  Cape  Cod,  but  we  find  no  actual  records  of  its  occurrence  farther  south  than 
Is  mentioned. 

'  Quoted  from  a  letter. 


Apparently  a  period  of  this  sort  occurred  about 
the  middle  of  the  past  century,  for  Perley,  writing 
in  1852,  reported  it  from  a  number  of  points  in  the 
neighborhood  of  St.  John,  New  Brunswick.  It 
seems  then  to  have  disappeared  from  the  Gulf  of 
Maine,  not  to  reappear  until  1903  when  it  was 
common  in  the  Bay  of  Fundy  in  May.  A  few 
were  again  taken  off  Passamaquoddy  Bay  in  that 
same  month  of  1915.8  And  this  was  the  prelude 
to  a  period  of  local  abundance,  for  capelin  were 
noticed  among  the  herring  in  the  weirs  of  the 
Passamaquoddy  Bay  region  in  October  1916, 
becoming  so  plentiful  by  the  end  of  November  that 
one  catch  of  3,000  pounds  of  fish  consisted  of  2,000 
pounds  of  capelin  and  only  1,000  of  herring. 
They  were  also  reported  at  various  localities  along 
the  New  Brunswick  coast  at  that  time.  Probably 
they  persisted  locally  in  the  Bay  of  Fundy  through- 
out the  winter  of  1916-1917,  for  large  numbers  of 
capelin  appeared  in  Minas  Basin  in  the  following 
May  and  June.  We  find  do  record  of  capelin 
within  the  limits  of  the  bay  in  1918,  but  they  were 
taken  again  in  1919  in  50  fathoms  of  water  off 
Passamaquoddy  Bay  in  January,  February,  and 
March,  and  they  appeared  with  smelts  a  month 
later  as  far  west  as  the  Penobscot  River,  pene- 
trating far  inland.  None,  however,  have  been  seen 
in  the  Gulf  of  Maine  since  then,  so  far  as  we  have 
been  able  to  learn. 

Smelt  Osmerus  mordax  (Mitchill)  1815 
Salt-water  smelt 

Jordan  and  Evermann,  1896-1900,  p.  523. 

Description. — The  smelt  is  distinguishable  from 
all  other  fish  common  in  our  waters  by  its  slender 
form,  combined  with  a  long  pointed  head,  large 
mouth,  a  small  but  evident  adipose  fin  standing 
above  the  rear  part  of  the  anal,  and  a  deeply 
forked  tail.  The  location  of  its  dorsal  fin  above 
the  ventrals  instead  of  in  front  of  them,  and  its 
much  larger  mouth  and  small  eye  separate  it  from 
the  argentine.  The  large,  fang-like  teeth  on  the 
smelt's  tongue,  its  larger  scales  (of  which  there 
are  about  75  along  each  row  on  the  sides,  all  alike 
in  the  two  sexes),  its  shorter  adipose  fin,  its  nar- 
rower pectoral  fins,  that  its  lower  jaw  projects 
only  slightly  beyond  the  upper  and  its  scales  slip 
off  very  easily,  obviate  any  danger  of  confusing 

•  Huntsman  (Contrib.  Canadian  Biol.,  (1921)  1922,  p.  50)  and  Kendall 
(Copula,  No.  42,  1917,  pp.  28-30;  and  Copela,  No.  73,  1919,  pp.  70-71)  give 
details. 


136 


FISHERY   BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


Figube  57.- — Smelt  (Osmerus  mordax),  adult,  Woods  Hole.     From  Jordan  and  Evermann.     Drawing  by  H.  L.  Todd. 


it  with  the  capelin.  The  body  of  the  smelt  is  only 
about  one-fifth  as  deep  as  long  (exclusive  of 
caudal  fin),  with  broadly  rounded  back  but  flat- 
tened enough  sidewise  to  be  egg-shaped  in  cross 
section.  It  is  deepest  about  its  mid-length,  taper- 
ing thence  toward  the  head  and  toward  the  tail  (at 
least  in  fat  fish),  whereas  the  capelin  is  of  nearly 
uniform  depth  from  gill  opening  to  anal  fin  (p.  134) . 
Its  mouth  gapes  back  of  the  eye. 

Printed  accounts  of  the  smelt  usually  credit  it 
with  a  peculiar  "cucumber"  odor,  and  smelt 
fishermen  often  speak  of  a  trace  of  this,  but  it  is  so 
faint  that  we  have  never  noticed  it  though  we  have 
caught  and  handled  many.9 

Color. — Transparent  olive  to  bottle  green  above, 
the  sides  are  of  paler  cast  of  the  same  hue  but  each 
with  a  broad  longitudinal  silvery  band.  The  belly 
is  silvery,  while  the  fins  and  body  are  more  or  less 
flecked  with  tiny  dusky  dots.  This  color  pattern 
is  shared  by  another  slender  little  fish,  the  silver- 
side  (p.  302),  but  the  latter  has  two  large  dorsal 
fins,  so  there  is  no  danger  of  confusing  the  smelt 
with  it. 

Size. — Smelt  grow  to  a  maximum  length  of 
about  13  or  14  inches.  Few,  however,  are  more 
than  a  foot  long,  and  adults  run  only  about  7  to 
9  inches.  Smelt  weigh  from  1  to  6  ounces  accord- 
ing to  size  and  fatness. 

Habits.10 —  The  smelt  is  an  inshore  fish,  con- 
fined to  so  narrow  a  zone  along  the  coast  that 
none  has  ever  been  reported  more  than  a  mile  or 
so  out  from  the  land,  or  more  than  two  or  three 
fathoms  in  depth,  while  many  spend  the  whole 
year  in  estuarine  situations. 

•  The  European  smelt  (0.  eptrlanus)  smells  so  strong  that  It  Is  not  held  In 
very  high  esteem  as  a  food-fish. 

i»  Kendall  (Bull.  V.  8.  Bur.  Fish.,  vol.  42, 1927,  p.  244)  has  given  a  detailed 
account  of  the  habits,  distribution,  and  catches  of  the  smelt  of  the  New 
England  coast,  also  of  the  landlocked  populations. 


Young  smelts  certainly,  and  old  ones  probably, 
travel  in  schools,  which  are  mostly  composed  of 
fish  of  a  size,  hence  probably  are  the  product  of 
one  year's  hatching,  and  they  five  pelagic,  not 
on  the  bottom,  though  confined  to  shoal  water. 

Most  authorities  describe  the  smelt  as  feeding 
on  small  crustaceans,  which  is  correct  as  far  as  it 
goes,  for  shrimp  (both  decapods  and  mysids) 
and  gammarids  are  probably  its  favorite  food,  and 
shrimp  were  long  considered  the  best  smelt  bait. 
But  it  has  been  found  that  pieces  of  "sea  worms" 
(Nereis)  are  more  attractive  to  the  larger  smelt 
(at  least  in  Massachusetts  Bay).  Small  fish  also 
form  an  important  item  in  the  diet.  We  have, 
for  example,  found  smelts  taken  in  the  Sheepscot 
River  in  May  packed  full  of  young  herring,  and 
have  caught  many  with  small  mnmmichogs 
(FundvJus)  as  bait;  while  dinners,  anchovies, 
launce,  sticklebacks,  silversides,  and  alewives  have 
been  identified  from  smelt  stomachs  at  Woods 
Hole.  The  Woods  Hole  diet  list  also  includes 
shellfish,  squid,  annelid  worms  (Nereis),  and  crabs, 
but  even  as  greedy  a  fish  as  the  smelt  ceases  to 
feed  during  its  spawning  visits  to  fresh  water. 
Young  smelt  depend  chiefly  on  copepods  and  on 
other  minute  pelagic  crustaceans.  Smelt  fisher- 
men are  familiar  with  the  fact  that  a  smelt  ap- 
proaches a  bait  slowly,  then  stops,  and  appears  to 
suck  it  in."  If  the  smelt  take  their  living  prey 
in  this  same  way,  it  is  somewhat  of  a  mystery 
how  they  succeed  in  capturing  animals  as  active 
as  shrimps  and  small  fish. 

Smelt,  like  alewives,  shad,  and  salmon,  make 
their  growth  in  salt  water,  but  run  up  into  fresh 
water  to  spawn. 

The  summer  habitat  of  the  smelt  varies  off 

H  This  method  of  feeding  seems  first  to  have  been  doscribed  In  print  by 
'•Orif'  (Forest  and  Stream,  vol.  54,  No.  8,  Feb.  24,  1900,  p.  151). 


FISHES   OF  THE  GULF  OF   MAINE 


137 


different  parts  of  the  coast  of  the  Gulf,  depending 
on  the  summer  temperature  of  the  water  and 
perhaps  on  the  food  supply.  Most  of  them  desert 
the  harbors  and  estuaries  of  the  Massachusetts 
Bay  region  and  along  the  southern  coast  of  Maine 
during  the  warmest  season.  But  it  is  probable 
that  they  move  out  only  far  enough  to  find  cooler 
water  at  a  slightly  greater  depth,  and  a  few  may 
be  found  in  harbors  through  the  summer.  Smelt, 
for  instance,  are  caught  then  in  Cohasset  Harbor 
in  some  years,  but  not  in  others ;  and  east  of  Penob- 
scot Bay,  where  the  surface  temperature  does  not 
rise  so  high  as  off  Massachusetts,  smelt  are  to 
be  found  in  the  harbors,  bays,  and  river  mouths 
all  summer,  and  are  sometimes  taken  in  numbers 
then  in  the  weirs.12 

Adult  smelt  gather  in  harbors  and  brackish 
estuaries  early  in  autumn,  where  smelt  fishing 
with  hook  and  line  is  in  full  swing  by  October. 
The  schools  then  tend  to  move  into  the  smaller 
harbors  on  the  flood  tide,  and  out  again  on  the 
ebb,  especially  if  the  tidal  current  is  strong,  as  it 
is  in  Cohasset,  a  locality  with  which  we  are  famil- 
iar. But  some  smelt  remain  over  the  ebb  in  the 
deeper  basins.  And  some  of  them  have  run  as 
far  as  the  head  of  tide  by  the  time  the  first  ice 
forms  in  December.  Most  of  them  winter  be- 
tween the  harbor  mouths  and  the  brackish  water 
farther  up;  the  maturing  fish  commence  their 
spawning  migration  into  fresh  water  as  early  in 
the  spring  as  the  ice  goes  out  of  the  streams  and 
the  water  warms  to  the  required  degree. 

Temperature  observations  by  the  Massachusetts 
Commission  show  that  the  first  smelt  appear  on 
the  spawning  beds  in  Weir  Eiver,  a  stream  empty- 
ing into  Boston  Harbor,  when  the  temperature  of 
the  water  rises  to  about  40-420.13  This  may  take 
place  as  early  as  the  first  week  in  March  or  as 
late  as  the  last,  about  Massachusetts  Bay,  depend- 
ing on  the  forwardness  of  the  season  and  on  the 
particular  stream.  The  chief  production  of  eggs 
takes  place  in  temperatures  of  50-57°,  and  spawn- 
ing is  completed  in  Massachusetts  waters  by  about 
the  10th  or  15th  of  May,  year  in  and  year  out. 
East  of  Portland,  smelt  seldom  commence  to  run 
before  April,  and  continue  through  May.  In  the 
colder  streams  on  the  southern  shores  of  the  Gulf 


of  St.  Lawrence  they  do  not  spawn  until  June. 
On  the  other  hand,  they  may  commence  spawn- 
ing as  early  as  February  along  the  southern  New 
England  coast  west  of  Cape  Cod. 

As  a  rule  smelt  do  not  journey  far  upstream; 
many,  indeed,  go  only  a  few  hundred  yards  above 
tidewater,  whether  the  stream  be  small  or  large. 
Thus  Dr.  Huntsman  informs  us  that  the  smelt  that 
enter  the  estuary  of  the  Stewiacke  River,  Nova 
Scotia  (a  tributary  of  the  lower  Shubenacadie, 
near  the  head  of  the  Bay  of  Fundy)  spawn  only 
in  the  tidal  part.  And  some  spawn  in  slightly 
brackish  water  in  certain  ponds  back  of  barrier 
beaches  (e.  g.,  Straits  Pond,  Cohasset,  Mass.). 
But  flooding  with  salt  water,  which  sometimes 
happens,  kills  the  eggs. 

The  adult  smelts  return  to  salt  water  immedi- 
ately after  spawning  to  spend  the  summer  either 
in  the  estuary  into  which  the  stream  in  which 
they  spawn  empties  or  in  the  sea  close  by.  On 
the  Massachusetts  coast  north  of  Cape  Cod  all 
the  spent  fish  have  left  fresh  water  by  the  middle 
of  May,  earlier  in  some  years.  On  the  Maine 
coast,  too,  a  good  proportion  of  the  spent  fish  are 
in  salt  water  by  the  first  weeks  in  May;  thus  we 
have  seen  a  bushel  of  large  smelt  taken  in  a  weir 
at  Cutler  (near  the  mouth  of  the  Grand  Manan 
Channel)  as  early  as  May  4. 

The  eggs  average  about  1.2  mm.  (0.05-inch)  in 
diameter  and  they  sink  to  the  bottom,  where 
they  stick  in  clusters  to  pebbles,  to  each  other,  or 
to  any  stick,  root,  grass,  or  water  weed  they  chance 
to  touch.  According  to  the  Manual  of  Fish 
Culture  a  female  weighing  as  little  as  2  ounces 
will  produce  between  40,000  and  50,000  eggs;14 
The  eggs  of  the  closely  allied  European  smelt 
{Osmerus  eperlanus)  hatch  in  8  to  27  days,  accord- 
ing to  temperature,  and  the  incubation  period  of 
the  American  fish  is  the  same,  probably,  for  smelt 
eggs  are  reported  as  hatching  in  13  days  at  the 
Palmer  (Mass.)  hatchery. 


"Atkins  (Fish.  Ind.  U.  8.,  sect.  5,  vol.  1,  1887,  pp.  690-693)  gives  much 
Information  on  the  smelt  In  Maine. 

»  Kendall  (Bull.  U.  S.  Bur.  Fish.,  vol.  42,  1927,  pp.  231-233)  summarizes 
these  observations  and  elves  additional  information  for  streams  on  tho  coast 
of  Maine. 


Figure  58. — Smelt  larva,  26  mm. 

The  smelt  has  proved  a  favorable  fish  for 
artificial  hatching  and  large  numbers  of  fry  are  so 
produced  yearly  in  Massachusetts,  the  eggs  being 

»  Kept.  U.  S.  Fish  Comm.,  1897,  p.  188. 


138 


FISHERY   BULLETIN    OF   THE    FISH   AND  WILDLIFE   SERVICE 


taken  in  Weir  Kiver,  just  mentioned,  and  it  has 
proved  possible  to  re-establish  smelt  by  intro- 
ducing the  eggs  or  fry  into  streams  from  which  it 
has  been  extirpated.  For  example,  good  smelt 
fishing  was  reported  in  "Poorhouse  Brook," 
Saugus,  a  tributary  of  Boston  Harbor,  three  years 
after  the  stream  was  stocked  with  eggs,  and 
attempts  have  been  similarly  successful  on  Long 
Island,  N.  Y.  Maintenance  of  the  stock  is  a 
question  either  of  providing  accessible  spawning 
grounds  of  sufficient  extent,  or  of  making  up  for 
lack  of  such  by  artificial  propagation. 

The  precise  season  when  young  smelt  go  down 
to  the  sea  in  the  Gulf  of  Maine  streams  is  yet  to 
be  learned;  probably  early  in  summer.  We 
seined  several  hundred  fry,  1  %  to  1  %  inches  long, 
October  1,  1924,  on  a  beach  of  Mount  Desert 
Island,  evidence  that  the  rate  of  growth  is  about 
the  same  for  our  smelt  during  its  first  summer  and 
autumn  as  for  the  European,  i.  e.,  to  a  length  of 
1%  to  2%  inches. 

Most  of  the  smelt  evidently  do  not  spawn  until 
they  have  passed  an  autumn,  a  winter,  a  summer, 
and  a  second  winter  in  salt  water. 

General  range. — East  coast  of  North  America 
from  eastern  Labrador,  Strait  of  Belle  Isle,  and 
the  Gulf  of  St.  Lawrence  southward  regularly  to 
New  Jersey,  and  reported  to  Virginia;  running  up 
streams  and  rivers  to  spawn.  Smelt,  also,  are 
landlocked  naturally  in  many  lakes  and  ponds  in 
New  Hampshire  and  in  Maine,  also  in  Lake 
Champlain,  and  in  various  Canadian  lakes.18 

Occurrence  in  the  Gulf  of  Maine. — The  smelt  is  a 
familiar  little  fish  around  the  entire  coast  of  the 
Gulf  of  Maine,  but  varies  greatly  in  abundance 
from  place  to  place  according  to  the  accessibility 
of  streams  suitable  for  spawning,  from  which  it 
seldom  wanders  far  alongshore.  Smelt  are  plenti- 
ful, still,  all  around  the  inner  parts  of  Massachu- 
setts Bay  and  its  tributary  harbors,  though  many 
of  the  local  streams  are  barred  to  them  now;  thence 
northward  and  eastward  all  along  the  coast  of 
Maine;  tolerably  so  in  the  region  of  Passama- 
quoddy  Bay  (catch  for  Charlotte  County,  New 
Brunswick,  7,400  pounds  in  1945),  and  more  so 
along  the  western  shore  of  Nova  Scotia  (60,100 
pounds  for  Yarmouth  County  in  1945).  But 
they  are  less  plentiful  passing  inward  along  the 
Nova  Scotia  shore  of  the  Bay  of  Fundy,  as  illus- 


'•  The  European  smelt  Is  landlocked  in  many  lakes  In  northern  Europe. 


trated  by  catches  in  1945  of  20,100  pounds  for 
Digby  County,  but  only  7,600  pounds  for  Kings 
County,  2,000  pounds  for  Hants,  and  1,800  for 
Colchester  (covering  the  Minas  Basin  region). 
So  few  smelt  exist  along  the  New  Brunswick  side 
of  the  Bay,  inward  from  the  Passamaquoddy 
region,  that  none  at  all  were  reported  for  that 
stretch  of  coast  in  any  year  during  the  period 
1939-1945.  Doubtless  this  scarcity  up  the  Bay 
is  "due  to  absence  of  streams  suitable  for  spawn- 
ing, and  the  general  turbidity  of  the  water,"  as 
Jeffers  has  remarked.1" 

Abundance. — Smelt  once  were  so  plentiful  in  the 
Back  Bay  at  Boston  (now  mostly  filled  in)  that 
"distinguished  merchants  of  lower  Beacon  Street 
might  be  seen,  at  early  hours,  eagerly  catching 
their  breakfast  from  their  back  doors."17  Those 
happy  days,  however,  are  long  since  past,  and 
smelt  certainly  are  not  so  numerous  as  they  were 
even  50  years  ago,18  around  the  Massachusetts 
shoreline  of  our  Gulf,  where  various  streams 
either  have  been  closed  to  them,  or  have  been 
rendered  uninhabitable  by  pollution.  But  enough 
still  remain  to  provide  sport  for  thousands  of 
anglers,19  and  we  still  hear  of  an  occasional  catch 
there  of  many  dozens  by  some  one  lucky  enough 
to  hit  a  run  of  fish  at  the  right  time  and  tide. 

In  1938,  when  a  special  effort  seems  to  have 
been  made  to  gather  smelt  statistics,  the  reported 
catch  for  the  inner  part  of  Massachusetts  Bay 
and  northward  to  the  New  Hampshire  line  was 
25,900  pounds,  or  some  300,000  fish,  if  they  ran 
about  a  dozen  to  the  pound.  The  yearly  catch 
reported  for  the  coast  of  Maine,  added  to  that  of 
the  Passamaquoddy  area  (which  form  one  faunal 
unit  so  far  as  the  smelt  is  concerned)  averaged 
about  644,000  pounds  during  the  period  1937  to 
1946,20  or  perhaps  some  8,000,000  fish;  about 
61,000  pounds  for  Digby  and  Yarmouth  Counties, 
Nova  Scotia,  combined,  which  covers  most  of  the 
catch  for  the  Gulf,  north  and  east  of  New  Hamp- 
shire. 

The  catches  of  smelt  that  are  made  along  the 
coasts  of  Maine,  New  Brunswick,  and  Nova 
Scotia  may  seem  impressive  if  taken  by  them- 

'•  Ann.  Rept.  Biol.  Board  Canada,  (1931)  1932,  p.  27. 

"  Mass.  Rept.  for  1870,  p.  23. 

»  Kendall  (Bull.,  U.  S.  Bur.  Fish.,  vol.  42,  1927,  pp.  244-249)  gives  many 
interesting  details  as  to  catches  in  Massachusetts. 

18  Smelt  fishing  has  long  been  restricted  to  hook  and  line  along  this  part 
of  the  coast. 

*>  Maximum  675,700  pounds  in  1945,  minimum  316,400  pounds  in  1939. 
No  data  are  available  for  Maine  for  the  years  1941  or  1942. 


FISHES   OF  THE  GULF  OF   MAINE 


139 


selves.  But  Miramichi  Bay,  alone,  on  the  south- 
ern shore  of  the  Gulf  of  St.  Lawrence  yields 
yearly  between  three  and  four  times  as  much 
smelt  as  does  the  entire  coastline  of  the  Gulf  of 
Maine.21 

Catch  records  do  not  suggest  any  striking 
alteration  in  the  abundance  of  smelts  during  the 
past  10  years  or  so  for  Maine  or  for  the  Canadian 
shores  of  the  Gulf.  But  they  seem  to  have  been 
somewhat  more  plentiful  along  the  Maine  coast 
previous  to  the  early  1900's,  for  catches  of 
1,125,268  to  1,279,550  pounds  there  in  1887,  1888, 
and  1902  have  not  been  equaled  since  then,  the 
nearest  approach  being  968,300  pounds  in  1945. 

We  are  often  asked  what  effect  the  disappear- 
ance of  the  eel  grass  (Zostera)  from  our  coasts  has 
had  on  the  abundance  of  the  smelt.  Unfor- 
tunately, the  statistics  of  the  yearly  catch  do  not 
yield  any  clear  answer.  Neither  can  we  offer  any 
convincing  explanation  for  the  violent  fluctuations 
that  take  place  from  year  to  year  in  the  abundance 
(or  availability?)  of  smelts  at  one  point  or  another. 
Fishermen  report,  for  example,  that  they  were  far 

»  Average  reported  catch  for  Northumberland  County    1937-1946,  wa» 
2.258,030  pounds. 


less  plentiful  in  Massachusetts  Bay  and  in  the 
Great  Bay  region,  N.  H.,  in  1950  than  they  were 
in  either  of  the  two  previous  years. 

The  smelt  also  has  a  great  recreational  value, 
smelt  fishing  being  a  favorite  pastime  for  home 
consumption.  As  many  as  2,326  people,  for  in- 
stance, have  been  counted  fishing  at  one  time  about 
Houghs  Neck  in  Boston  Harbor,  and  this  same 
sort  of  thing  is  to  be  seen  up  and  down  the  Massa- 
chusetts coast  in  harbors  and  stream  mouths  in 
autumn.  Many  smelt  are  caught  in  Great  Bay, 
N.  H.,  in  good  years,  through  the  ice  for  the  most 
part.  And  this  applies  equally  to  many  localities 
along  the  coast  of  Maine.  So  plentiful  are  the 
fish  on  occasion  and  so  greedily  do  they  bite, 
especially  on  the  flood  tide,  that  it  is  usual  to 
number  the  catch  about  Massachusetts  Bay  by 
the  dozens  rather  than  by  the  individual  fish. 
Sea  worms  (Nereis)  are  generally  considered  the 
best  bait,  especially  for  the  larger  smelt,  shrimp 
the  second  best,  small  minnows  or  clams  a  poor 
third.  Smelt  have  also  been  taken  with  a  small 
red  artificial  fly  in  the  Gulf  of  St.  Lawrence,  and 
perhaps  elsewhere. 


THE  ARGENTINES 
FAMILY  ARGENTINIDAE 


The  argentines  resemble  the  smelts  in  most  of 
their  external  characters.  But  their  mouths  are 
much  smaller,  with  the  upper  jawbone  reaching 
back  only  about  even  with  the  front  of  the  eye, 
and  the  entire  base  of  their  rayed  dorsal  fin  is  in 
front  of  the  ventral  fins. 

Argentine  Argentina  silus  Ascanius  1763 
Herring  smelt 

Jordan  and  Evermann,  1896-1900,  p.  526. 

Description. — The  argentine  has  a  pointed  nose, 
deeply  forked  tail,  and  slender,  compressed  body, 
but  it  has  much  larger  eyes  than  either  smelt  or 
capelin,  a  character  no  doubt  associated  with  its 
deep-water  home;  its  mouth  is  much  smaller,  not 
gaping  back  even  as  far  as  the  eye;  and  its  dorsal 
fin  stands  wholly  in  front  of  the  ventrals,  instead 
of  above  them  as  it  does  in  both  the  smelt  and  the 
capelin.22 

"  The  anatomy  of  Argentina  silus  Is  described,  and  records  along  the 
American  coast  are  given  by  Kendall  and  Crawford  (Jour.  Washington 
Acad.  Sol.,  vol.  12,  No.  1,  January  1922,  pp.  8-19). 


The  body  of  the  argentine  (about  one-fifth  as 
deep  as  long)  tapers  toward  both  head  and  tail,  but 
its  sides  are  so  flat,  and  its  back  and  belly  so 
broad,  that  it  is  nearly  rectangular  in  cross  section 
instead  of  oval.  Its  scales,  too,  are  larger  than 
those  of  the  smelt,  there  being  only  60  to  70  rows 
along  the  lateral  line.  Its  adipose  fin  is  very  small 
and  its  jaws  are  toothless,  though  its  palate  and 
tongue  are  armed  with  small  teeth. 

Color. — The  color  of  the  adult  is  variously  de- 
scribed by  different  authors.  All  agree,  however, 
that  the  back  is  brownish  or  olivaceous,  the  sides 
silvery  or  with  iridescent  golden  or  brassy  luster, 
and  the  belly  white.     The  adipose  fin  is  yellowish. 

Size. — The  argeutine  is  a  larger  fish  than  the 
smelt  or  the  capelin,  growing  to  a  length  of  about 
18  inches. 

Habits. — Nothing  is  known  of  the  life  of  the 
herring  smelt  in  our  Gulf,  and  little  enough  is 
known  of  it  in  Scandinavian  waters,  where  it  is 
sometimes  caught  on  deep  set-lines  baited  with 
herring  or  mussels,  and  where  it  is  occasionally 


140 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  59. — Argentine  (Argentina  silus).     A,  adult,  Biddeford  Pool,  Maine;  from  Goode  and  Bean,  drawing  by  H.  L. 
Todd;  B,  egg;  C,  larva,  28  mm.;  D,  larva,  45  mm.     B-D,  European,  after  Schmidt. 


swept  up  to  the  surface  like  other  deep-sea  fishes 
by  some  upwelling  of  the  water,  to  drift  there 
helplessly.  Its  eggs  float  chiefly  in  the  deeper 
water  layers,  seldom  rising  to  the  surface,  and 
they  are  among  the  largest  of  buoyant  fish  eggs 
(3  to  3.5  mm.  in  diameter),  with  flat  oil  globule 
(0.95  to  1.16  mm.)  and  vacuolated  yolk.  Newly 
hatched  larvae  are  about  7.5  mm.  long  and  have 
a  large  yolk  sac,  but  this  has  been  absorbed  when 
they  have  grown  to  a  length  of  12  mm.  and  a  line 
of  spots  has  appeared  along  the  belly.  The  fin 
rays  are  formed  by  the  time  the  little  fish  has 
reached  45  mm.,  the  anus  has  moved  forward,  and 
the  forked  outline  of  the  tail  is  apparent,  but  the 
ventral  fins  do  not  appear  untd  the  larva  is  about 
50  mm.  long. 

General  range.— North  Atlantic,  usually  in  water 
as  deep  as  80  to  300  fathoms ;  known  from  northern 
Norway  south  to  the  northern  part  of  the  North 
Sea  on  the  European  side,  from  the  Nova  Scotia 
Banks  to  the  offing  of  southern  New  England  on 
the  American  side.23 

Occurrence  in  the  Gulf  of  Maine. — The  argentine 
was  considered  rare  in  our  waters  until  recent- 
ly.    Some  specimens  have  been  brought  in  from 

»  For  recent  records  of  argentlnes  off  Nova  Scotia,  see  McKenzle  and 
Homans,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  19,  No.  3,  1938,  p.  277  and 
McKenzle,  Proc.  Nova  Scotia  Inst.  Scl.,  20, 1939,  p.  16. 


widely  scattered  localities  around  the*  coast,  name- 
ly, Belfast,  Biddeford  Pool,  and  Fletchers  Neck, 
Maine;  and  from  Hampton  Beach,  N.  H.  It  has 
proved,  with  the  development  of  otter  trawling, 
that  argentines  are  fairly  common  all  around  the 
edges  of  Georges  Bank  and  off  Cape  Cod  in  mod- 
erately deep  water.  It  is  not  unusual  for  one  haul 
of  the  trawl  to  bring  in  from  one  to  a  dozen  from 
depths  of  30  to  100  fathoms,  with  much  larger 
numbers  taken  occasionally;  one  vessel,  for  exam- 
ple, trawled  15,000  pounds  on  the  northeastern 
edge  of  Georges  Bank  in  about  100  fathoms  during 
a  week  in  mid-September  1929.  Evidently  there 
are  at  least  a  few  argentines  in  the  deep  trough  of 
the  Gulf  also.  Firth24  reports  that  ten  were  taken 
at  90  fathoms  on  the  northwestern  slope  of  Georges 
Bank  on  June  18;  and  the  Albatross  II  trawled 
one  at  1 15  fathoms  off  Mount  Desert  Rock.  They 
spawn  to  some  extent  in  the  Gulf,  for  on  April  17, 
1920,  a  townet  haul  on  the  Albatross  I  from  109 
fathoms  in  the  southeastern  part  of  the  Gulf  basin 
yielded  43  eggs,  unmistakably  of  argentine  par- 
entage, while  we  have  taken  a  scattering  of  argen- 
tine fry  at  localities  as  widely  separated  as  the 
offing  of  Mount  Desert  Rock  and  the  northwestern 
edge  of  Browns  Bank. 

»  Firth.  Bull.  Boston  Soc.  Nat.  Hist.,  61. 1940,  p.  10. 


FISHES   OF  THE  GULF  OF   MAINE 

LUMINESCENT  FISHES 


141 


FAMILIES    MYCTOPHIDAE,   MAUROLICIDAE,  CHAULIODONTIDAE,   GONOSTOMIDAE, 

STOMIATIDAE,  AND  STERNOPTYCHIDAE 


These  families  include  a  heterogeneous  assem- 
blage of  small  oceanic  fishes,  that  are  primitive  in 
some  respects,  but  are  highly  specialized  in  others 
for  existence  in  mid-depths,   on  the  high  seas. 


They  all  have  light-producing  organs,  which  no 
other  Gulf  of  Maine  fish  has;  this  is  the  only  reason 
why  we  group  them  together  here. 


KEY  TO  GULF  OF  MAINE  LUMINESCENT  FISHES 

1.  Trunk  at  least  Yi  as  deep  as  it  is  long  from  tip  of  snout  to  base  of  tail  fin;  front  part  of  rayed  dorsal  fin  is  a 

hard  triangular  plate,  supported  by  7  or  8  spines Hatchet  fish,  p.  149 

Trunk  less  than  %  as  deep  as  it  is  long  from  tip  of  snout  to  base  of  tail  fin;  rayed  dorsal  fin  does  not  commence  with  a 
hard  plate  or  hard  spines 2 

2.  Mouth  does  not  gape  back  as  far  as  the  eye Pearl  sides,  p.  144 

Mouth  gapes  back  beyond  the  eye 3 

3.  No  barbel  on  the  chin;  the  ventral  fins  are  about  mid- way  of  the  trunk;  origin  of  rayed  dorsal  fin  either  in  front  of  mid- 

length  of  trunk  or  at  least  not  much  behind  it 4 

There  is  a  long  fleshy  barbel  on  the  chin;  the  ventral  fins  are  considerably  behind  the  mid-length  of  the  trunk;  the 
rayed  dorsal  fin  is  far  back,  close  to  the  tail  fin 7 

4.  The    rayed    dorsal    fin    is    far   in    advance    of   the    ventrals;    the    jaws    are    armed    with    long    and    conspicuous 

fangs Viperfish,  p.  145 

The  rayed  dorsal  fin  is  about  over  the  ventral  fins  (it  may  be  a  little  in  front  of  them  or  a  little  behind) ;  the  teeth  are 
small 5 

5.  Eyes    very   small;    no   adipose   fin    behind   the   rayed   dorsal   fin;    anal   fin    reaches   nearly   to    the    base    of   the 

caudal Cyclothone,  p.  146 

Eyes  very  large;  there  is  an  adipose  fin  behind  the  rayed  dorsal;  there  is  a  considerable  interspace  between  the  rear  end 
of  the  anal  fin  and  the  origin  of  the  tail  fin 6 

6.  There  are  3  or  4  separate  luminescent  dots  at  the  base  of  the  caudal  fin;  the  Gulf  of  Maine  species  has  a  large  lumines- 

cent patch  on  the  snout Headlight  fish,  p.  142 

There  are  only  2  separate  luminous  dots  at  the  base  of  the  caudal  fin;  the  snout  does  not  have  a  large  luminescent 
patch Lanternfish,  p.  143 

7.  The  point  of  origin  of  the  anal  fin  is  in  advance  of  the  origin  of  the  rayed  dorsal  fin  by  a  distance  about  as  long  as 

the    diameter    of    the    eye;    the    tip    of    the    chin    barbel    is    distinctly    swollen    as     well    as    bearing    several 

filaments Stomioides,  p.  147 

The  point  of  origin  of  the  anal  fin  is  not  in  advance  of  the  rayed  dorsal  fin,  the  tip  of  the  chin  barbel  is  not  swollen..     8 

8.  Each  side  has  only  about  68  luminescent  spots;  there  is  a  large  luminescent  patch  crossing  the  top  of  the  cheek,  behind 

the  eye;  the  point  of  origin  of  rayed  dorsal  fin  is  in  advance  of  origin  of  anal  fin  by  a  distance  about  as  long  as  the 
diameter  of  the  eye;  the  tip  of  the  lower  jaw  does  not  enclose   the   tip   of   the    upper   jaw   when   the    mouth 

is  closed Trigonolampa,  p.  148 

Each  side  has  about  85  luminescent  spots;  the  side  of  the  cheek  behind  the  eye  does  not  have  a  large  luminescent 
patch;  the  point  of  origin  of  rayed  dorsal  fin  is  about  over  origin  of  anal  fin;  the  tip  of  the  lower  jaw  encloses  the  tip 
of  the  upper  jaw  when  the  mouth  is  closed Stomias,  p.  147 

LANTERN  FISHES.     FAMILY  MYCTOPHIDAE 


The  most  distinctive  external  characters  of  the 
lanternfishes  are  their  large  eyes  (situated  close  to 
the  tip  of  the  blunt  snout),  wide  mouths  gaping 
back  beyond  the  eye,  one  soft-rayed  dorsal  fin,  a 
deeply  forked  tail,  and  the  presence  of  a  series  of 
luminous  organs  as  conspicuous  pale  spots  along 
the  sides.  Some  of  them  have  an  adipose  fin  on 
the  back  behind  the  dorsal  fin,  but  others  lack 
this.  When  present,  this  fin  is  so  small  and 
fragile  that  it  is  apt  to  be  destroyed  by  the  rough 


treatment  the  fish  receive  in  the  tow  net  in  which 
they  are  taken.  They  most  aearly  resemble  the 
anchovy  (p.  118),  the  pearlsides  (p.  144),  and  the 
cyclothone  (p.  146)  among  Gulf  of  Maine  fishes; 
but  they  are  readily  distinguished  from  the  first 
of  these  by  the  presence  of  luminous  organs  and 
by  the  fact  that  the  snout  does  not  project 
beyond  the  mouth;  from  the  second  by  their 
much  wider  mouths;  and  from  the  third  by  their 
much  larger  eyes. 


142 


FISHERY  BULLETIN    OF  THE    FISH   AND   "WILDLIFE   SERVICE 


They  are  among  the  most  numerous  fishes  on 
the  high  seas,  where  they  live  at  a  considerable 
depth  by  day  but  often  rise  to  the  surface  at  night. 
Only  two  species  of  the  group,  representing  as 
many  genera  (Diaphus  and  Myctophum),  have 
been  recorded  within  the  Gulf  of  Maine.  But 
each  of  these  genera  includes  a  considerable 
number  of  species  that  are  common  along  the 
continental  slope  abreast  of  the  Gulf,  hence  are 
as  likely  to  stray  into  the  latter  as  are  the  two 
that  have  actually  been  found  there.  And  this 
applies  equally  to  various  other  genera  of  lantern- 
fishes. 

The  species  of  Diaphus  and  of  Myctophum  all 
resemble  one  another  in  general  appearance,  in 
having  a  short  dorsal  fin,  with  an  adipose  fin 
behiod  it;  a  deeply  forked  tail;  large  eyes;  wide, 
oblique  mouth;  and  numerous  luminous  organs 
along  the  sides;  all,  too,  are  blackishsilvery 
in  color.  The  members  of  each  genus  are 
separable  only  by  differences  in  the  arrangement 
of  the  luminous  organs.  Hence,  positive  identi- 
fication of  a  given  specimen  calls  for  the  services 
of  a  specialist  in  the  group.  Should  a  lanternfish 
be  taken  in  the  Gulf  in  which  tbe  arrangement  of 
luminous  organs  does  not  agree  precisely  with  the 
two  described  here,  we  suggest  that  it  be  sub- 
mitted to  the  U.  S.  Fish  and  Wildlife  Service  to 
be  named.26 


«  Parr  (Bull.  Bingham  Oceanog.  Coll.,  vol.  3,  art.  3,  1928),  and  Tanlng 
(Vldensk.  Meddel.,  Dansk  Naturhlst.  Forenlng,  vol.  86,  p.  49,  1928)  have 
recently  published  critical  synopses  of  the  lantemflshes. 


Headlight   fish   Diaphus   effulgens    (Goode   and 
Bean)  1895 

Jordan  and  Evermann,  1896-1900,  p.  566. 

Description. — This  curious  little  fish  is  separable 
from  the  lanternfish  (p.  143)  and  from  the  pearlsides 
(p.  144)  at  a  glance,  by  the  large  and  very  noticeable 
luminescent  patch  that  covers  the  entire  tip  of  its 
snout  (including  the  anterior  margin  of  the  orbit) 
and  that  extends  down  over  the  edge  of  the  upper 
jaw,  a  structure  that  has  no  parallel  in  any  other 
fish  regularly  inhabiting  the  Gulf  of  Maine.  It 
also  differs  from  the  pearlsides  in  its  much  more 
deeply  cleft  mouth,  its  even  larger  eyes,  in  the 
more  convex  dorsal  profile  of  its  head,  and  in  lack- 
ing the  regular  horizontal  row  of  luminescent 
spots  along  each  side  about  at  the  level  of  the 
pectoral  fin,  that  are  conspicuous  on  the  pearl- 
sides.2* 

The  four  separate  luminescent  spots  at  the  base 
of  the  tail  (besides  the  organ  on  its  snout)  separate 
it  from  its  close  relatives  of  the  genus  Myctophum 
(p.  143).  The  arrangement  of  the  fins  (all  of 
which  are  soft,  the  dorsal  with  about  15  rays, 
the  anal  with  about  16),  is  essentially  the  same 
as  in  the  latter,  and  in  the  pearlsides;  the  caudal 
fin  is  more  deeply  forked  than  in  the  pearlsides, 
the  adipose  fin  proportionately  shorter. 

Color. — The  color  has  not  been  described. 
Probably  it  is  black,  overlaid  more  or  less  with 

»  The  structures  along  the  lateral  line  shown  here  on  the  Illustration  of 
the  headlight  fish  are  large  scales,  not  luminescent  organs. 


Figure  60. — Headlight  fish  (Diaphus  effulgens),  Browns  Bank.     From  Goode  and  Bean.     Drawing  by  A.  H.  Baldwin. 


FISHES  OF  THE  GULF  OF   MAINE 


143 


silver,  with  the  luminescent  organs  pale  blue 
or  green. 

Size. — The  specimens  from  which  this  species 
was  originally  described  seem  to  have  been  about 
7  inches  long.27 

General  range  and  occurrence  in  the  Gulf  of 
Maine. — This  oceanic  species  is  only  a  stray 
within  the  limits  of  the  Gulf.  One  specimen  has 
been  found  in  the  stomach  of  a  cod  caught  on 
Browns  Bank,28  and  another,  also  from  a  cod 
stomach,  has  been  reported  on  Western  Bank  off 
the  outer  coast  of  Nova  Scotia.28 

Lanternfish  Myctophum  affine  (Liitken)  1892 

Jordan  and  Evermann,  1896-1900,  p.  570. 

Description. — The  most  noticeable  features  of 
this  little  oceanic  fish  are  its  silvery  black  color, 
the  luminous  dots  along  its  sides,  its  enormous 
eye  situated  close  to  the  tip  of  the  snout,  its  very 
deep  oblique  mouth,  and  its  deeply  forked  tail. 
The  anal  fin  is  mostly  or  wholly  behind  the  short, 
soft  dorsal,  acid  there  is  an  adipose  fin  behind  the 
latter,  as  in  the  headlightfish  (p.  142).  The  longer 
snout  and  smaller  mouth  of  Myctophum,  with  the 
fact  that  the  luminous  organs  on  its  snout  are  in 
the  form  of  small  dots  instead  of  a  large  patch 


covering  the  entire  tip  of  the  snout,  are'the  readiest 
field  marks  to  distinguish  it  from  the  latter.  The 
dorsal  profile  of  the  head  is  much  arched,  the 
body  moderately  flattened  sidewise,  tapering 
gently  backward  to  the  rather  deep  caudal  pe- 
duncle. The  location  of  the  luminescent  spots  is 
shown  in  the  drawing  (fig.  61). 

Color. — This  lanternfish  is  silvery  when  alive, 
the  silver  underlain  on  the  back  with  deep  brown- 
ish black,  the  sides  below  the  lateral  line,  and  the 
belly  varying  (below  the  silver)  from  dark 
brown  to  dusky  gray,  or  even  to  white  finely 
dotted  with  gray.  The  luminescent  organs  are 
pale  green  or  blue. 

Size. — All  members  of  the  genus  Myctophum 
are  small;  a  little  more  than  3K  inches  (89  mm.) 
is  the  maximum  length  recorded  for  this  particular 
species. 

General  range. — All  the  species  of  this  genus 
are  oceanic,  occurring  only  as  strays  inside  the 
edge  of  the  continent. 

Occurrence  in  the  Gulf  of  Maine. — Goode  and 
Bean30  report  the  capture  of  this  lanternfish  over 
the  southeast  slope  of  Browns  Bank  (lat.  42°  21' 
N.,  long.  65°  07'  W.)  at  104  fathoms,  which  still 
remains  the  only  record  for  it  in  the  Gulf  of 
Maine,31  or  for  any  Myctophum  for  that  matter. 


Figure  61. — Lanternfish  (Myctophum  affine).     After  Parr. 


PEARLSIDES.      FAMILY  MAUROLICIDAE 


The  Pearlsides  resembles  the  lanternfishes  (p. 
141)  in  shape  of  body,  but  it  has  a  shorter  rayed 


"  The  illustration  (Ooode  and  Bean,  Smithsonian  Contrlb.  Knowl.,  vol. 
31, 1895,  fig.  103),  about  6  Inches  long.  Is  characterized  In  the  legend  as  "slightly 
reduced." 

"  Reported  by  Goode  and  Bean  (Smithsonian  Contrlb.  Knowl.,  vol.  30, 
1895,  p.  88)  as  Aethoprora  effulgent. 

»  Vladykov,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  19, 1935,  p.  2. 


dorsal  fin,  a  longer  adipose  fin,  a  longer  anal,  and 
a  much  smaller  mouth. 

»  Smithsonian  Contrlb.  Knowl.  (vol.  30, 1895,  p.  72)  as  M.  opallnum. 

»>  It  Is  likely  that  Myctophum  alaciale  will  be  found  In  the  Oulf  of  Maine 
sooner  or  later,  judging  from  Its  widespread  distribution  In  the  boreal  belt 
of  the  Atlantic  and  from  the  fact  that  It  has  often  been  caught  at  the  surface. 
It  resembles  M.  affine  very  closely  In  appearance,  and  In  the  general  arrange- 
ment  of  the  luminous  organs,  but  d  InYrs  from  It  In  that  one  of  the  luminescent 
spots  above  the  base  of  the  ventral  fin  Is  elevated  above  the  others. 


144 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Pearlsides  Maurolicus  pennanti  (Walbaum)  1792 

Pearlfish 

Jordan  and  Everman,  1896-1900,  P.  577. 

Description. — The  presence  of  an  adipose  fin  be- 
tween the  dorsal  and  caudal  fins,  together  with 
luminous  organs,  distinguishes  the  pearlsides  from 
all  other  fishes  that  occur  regularly  in  the  Gulf  of 
Maine.  It  agrees  in  both  these  respects  with  the 
lanternfish  (p.  143)  and  with  the  headlightfish  (p. 
142),  but  it  has  a  much  smaller  mouth  and  a  longer 
adipose  fin  than  the  first  of  these,  and  it  lacks  the 
large  luminous  patch  on  the  snout  that  is  so  strik- 
ing a  feature  of  the  second.  Also,  the  pearlsides, 
with  its  herring-like  coloration  (p.  88)  differs  strik- 
ingly from  the  lanternfish,  which  has  a  black  back 
overlaid  with  silver;  and  probably  the  headlight 
fish  as  well. 


Figure    62. — Pearlsides   (Maurolicus    pennanti).      After 
Smitt. 

The  pearlsides  is  a  flat-sided,  large-headed  little 
fish,  its  body  (about  one-fifth  as  deep  as  long,  ex- 
cluding caudal  fin)  deepest  forward  of  the  ventral 
and  dorsal  fins;  its  eye  very  large;  its  lower  jaw  pro- 
jecting; its  mouth  oblique;  and  both  its  jaws  armed 
with  minute  teeth.  The  dorsal  fin  (about  1 1  or  12 
rays)  stands  above  the  space  between  the  ventrals 
and  the  anal;  the  anal  is  longer  than  the  dorsal. 
The  adipose  fin  (both  of  Woods  Hole  32  and  of 
Norwegian  33  examples)  is  low  and  long,  much  as  it 
is  in  the  capelin.34  The  caudal  fin  is  broad  and 
slightly  forked. 

The  pearlsides  has  been  described  as  without 
scales,  but  this  is  not  correct,  for  both  Scandinav- 
ian and  Woods  Hole  specimens  have  been  found  to 
be  clothed  with  large  but  extremely  thin  trans- 
parent scales.     There  is  no  definite  lateral  line. 


»  Sumner,  Osburn,  and  Cole,  Bull.  U.  S.  Bur.  Fish.,  vol.  31,  Pt.  2,  1913, 
p.  743. 

"  Smltt,  Scandinavian  Fishes,  vol.  1, 1892,  p.  933,  pi.  44,  flg.  3. 

"  Ooode  and  Bean  (Smithsonian  Contrlb.  Knowl.,  vol.  30,  1895  p.  96) 
describe  It  as  "very  small,"  but  probably  their  specimens  were  damaged. 


The  most  interesting  and  diagnostic  feature  of 
the  pearlsides  is  the  presence  of  a  series  of  lumi- 
nescent dots  situated  as  follows: 35  First,  12  pairs 
along  the  belly  between  the  pectoral  and  the  ven- 
tral fins,  followed  by  5  or  6  from  the  ventral  fins  to 
the  anal  fin,  and,  after  a  gap,  by  24  or  25  between 
the  center  of  the  anal  fin  and  the  base  of  the  caudal 
fin;  all  these  together  form  a  practically  continuous 
row  on  each  side  of  the  belly  from  throat  to  tail. 
Second,  there  is  a  row  of  larger  spots  a  little  higher 
up  on  each  side,  6  from  chin  to  pectoral  fin,  and  9 
thence  backward  to  the  ventrals.  Third,  there  is 
a  group  of  6  low  down  on  each  side  of  the  cheek  and 
throat;  there  is  likewise  a  spot  in  front  of  the  base 
of  each  pectoral  fin  and  2  on  the  chin. 

Color. — The  pearlsides  is  colored  much  like  a 
herring,  with  dark  bluish  or  greenish  back  and  lus- 
trous silvery-white  sides  and  belly.  The  lumines- 
cent spots  are  described  as  black  rimmed,  their 
centers  as  pale  blue  in  life  but  turning  yellow  in 
alcohol;  and  there  is  a  narrow  black  band  along  the 
base  of  the  anal  fin  and  from  there  to  the  base  of 
the  caudal,  the  latter  being  barred  with  a  similar 
black  band. 

Size. — Only  1  to  2%.  inches  long. 

Habits. — The  relatives  of  the  pearlsides  are  oce- 
anic, living  in  the  mid-depths  mostly  below  150 
fathoms,  but  the  pearlsides  itself  has  been  found  so 
often  in  the  stomachs  of  cod  and  of  herring  (fish 
that  do  not  descend  to  any  great  depth)  that  there 
is  no  reason  to  regard  it  as  a  "deep-sea"  stray,  nor 
has  it  ever  been  taken  far  from  land  so  far  as  we  can 
learn.  It  probably  spawns  in  early  spring,  females 
with  large  eggs  having  been  taken  in  Scottish 
waters  in  winter. 

General  range. — The  pearlsides  (there  are  several 
other  species  closely  allied  to  it)  ranges  widely  in 
the  open  Atlantic,  occurring  at  times  in  shoals  on 
the  coasts  of  Norway  and  in  British  waters.  It  is 
especially  common  off  the  coast  of  Scotland,  but 
has  not  been  recorded  often  on  the  American  side 
of  the  Atlantic. 

Occurrence  in  the  Gulf  of  Maine. — The  known 
occurrences  of  the  pearlsides  in  the  Gulf  have  been 
few.  Storer 36  (1867)  records  one  found  alive  on 
the  beach  at  Nahant,  Mass.,  in  December,  1837; 
another   taken   from   the   stomach   of  a   cod   at 


»  This  account  Is  based  chiefly  on  Smitt's  description  and  plate,  the  speci- 
mens we  have  seen  being  In  poor  condition. 
«  Fishes  of  Mass.,  1887,  p.  160,  as  Scopelus  humboldtil. 


FISHES   OF  THE  GULF  OF   MAINE 


145 


Provincetown;  a  third  picked  up  alive  there  in 
July,  1865  (pictured  by  Storer  on  pi.  25,  fig.  5); 
and  five  others  found  on  the  Provincetown  beach 
soon  afterward.  We  have  seen  one  specimen  41 
mm.  long  taken  from  the  stomach  of  a  cod,  on 
Platts  Bank,  July  27,  1924;  one  43  mm.  long,  also 
from  a  cod's  stomach,  on  Cashes  Ledge,  August 
16,  1928;  and  four,  32  to  39  mm.  long,  taken  from 
the  stomachs  of  <two  pollock  that  we  caught  in  20 
fathoms,  7  miles  southeast  of  Bakers  Island, 
Mount  Desert,  Maine,  July  24,  1930.  It  has 
been  found  twice  at  Grand  Manan,37  and  speci- 
mens were  picked  up  on  the  beach  at  Campobello 
Island  at  the  mouth  of  the  Bay  of  Fundy  in  July 


1914,38  while  others  were  taken  from  the  stomach 
of  a  pollock  caught  near  by.  It  has  also  been 
recorded  twice  near  Woods  Hole. 

These  locality  records  are  distributed  widely 
enough  to  show  that  it  is  to  be  expected  anywhere 
in  our  Gulf.  And  we  suspect  that  the  pearlsides 
is  not  as  scarce  there  as  the  paucity  of  actual 
records  for  it  might  suggest  (in  fact,  Storer  tells 
us  that  a  Nahant  fisherman  reported  finding  them 
repeatedly  in  the  stomachs  of  haddock  many 
years  ago) ,  but  that  it  keeps  out  of  sight,  being  an 
inhabitant  of  the  deeper  water  layers  as  its 
luminescent  organs  would  suggest,  coming  up  to 
the  surface  chiefly  at  night. 


VIPER  FISHES.     FAMILY  CHAULIODONTIDAE 


The  viper  fishes  have  slender  bodies,  bulldog- 
like faces  with  long  fangs;  the  first  dorsal  very  far 
forward,  the  anal  far  back;  and  no  barbel  on  the 
chin. 

Viperfish  Chavliodus  sloani  Bloch  and  Schneider 
1801 

Jordan  and  Evermann,  1896-1900,  p.  585. 

Description. — The  viperfish  not  only  has  lum- 
inescent organs,  but  it  is  very  different  in  general 
appearance  from  all  the  fishes  that  are  regular 
inhabitants  of  the  Gulf  of  Maine.  Most  obvious 
of  its  characteristics  is  its  bulldog-like  mouth.  It 
shares  this  with  its  fellow  strays,  Stomias  (p.  147), 
Stomioides  (p.  147)  and  Trigonolampa  (p.  148)  and 
the  general  form  is  much  alike  in  the  three.  But 
there  is  no  danger  of  confusing  it  with  any  one  of 
these  if  one  looks  closely,  for  the  viperfish  has  an 


»  Coi  (Bull.  Nat.  Hist.  Soc.  New  Brunswick,  14,  1896,  append.,  p.  65) 
reported  one  found  dead  there,  on  the  shore, 
a  Huntsman  (Contrlb.  Canadian  Biol.,  (1921)  1922,  p.  61.) 


adipose  fin  and  its  rayed  dorsal  fin  is  far  forward, 
whereas  Stomias,  Stomioides,  and  Trigonolampa 
have  no  adipose  fin  and  their  rayed  dorsal  fin 
stands  far  rearward. 

In  the  viperfish  the  lower  jaw  is  longer  than  the 
upper,  the  upper  is  armed  with  four  long  fangs  on 
each  side,  while  the  lower  has  a  series  of  pointed 
teeth  set  far  apart,  those  in  front  very  elongate 
and  all  of  them  so  long  that  tbey  project  when  the 
mouth  is  closed.  Furthermore,  the  snout  is  so 
short  that  the  very  wide  mouth  gapes  far  back  of 
the  eye.  The  body  is  about  seven  times  as  long 
as  deep,  flattened  sidewise,  deepest  close  behind 
the  head,  and  tapering  evenly  to  the  tail.  The 
very  short  dorsal  fin  (6  or  7  rays)  stands  far 
forward  and  its  first  ray  is  separate,  very  slender, 
and  about  half  as  long  as  the  fish  when  not  broken 
off,  as  it  usually  is.  The  ventrals  are  about 
midway  between  the  snout  and  the  origin  of  the 
anal  fin,  variously  pictured  as  either  larger  or 
smaller  than  the  dorsal.  The  small  anal  is  close 
to  the  caudal,  with  the  adipose  fin  over  it.     The 


Figure  63. — Viperfish  (Chauliodus  sloani),  southern  slope  of  Browns  Bank.     After  Goode  and  Bean. 


146 


FISHERY  BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


body  is  clothed  with  large  but  very  thin  scales. 
There  are  several  longitudinal  rows  of  small 
luminescent  spots  on  the  ventral  surface,  running 
from  throat  to  tail;  several  more  such  spots  on 
each  side  of  the  head ;  and  many  tiny  unpigmented 
dots  scattered  over  the  trunk.3' 

Color. — Greenish  above,  the  sides  with  metallic 
gloss;  blackish  below. 

Size. — Up  to  about  one  foot  long. 

Habits. — Nothing  is  known  of  its  habits  except 
that  it  is  an  inhabitant  of  the  mid-depths  of 
the  Atlantic  Basin  and  that  it  probably  does  not 
rise  closer  to  the  surface  than  150  or  200  fathoms 


except,  perhaps,  during  its  larval  stages.  Its 
teeth  suggest  a  rapacious  habit  but  there  is  no 
actual   record   of  its   diet. 

Occurrence  in  the  Gulj  of  Maine. — The  only 
definite  Gulf  of  Maine  records  are  of  one  specimen 
found  in  the  stomach  of  a  cod  caught  on  Georges 
Bank  in  1874,  and  of  a  second  found  in  the 
stomach  of  a  swordfish  that  was  harpooned  in 
the  gully  between  Browns  and  Georges  Banks  in 
1931.40  But  the  viperfish  may  be  expected  on 
the  offshore  banks  as  a  stray  at  any  time,  for 
several  have  been  taken  off  the  continental  slope 
abreast  of  southern  New  England  "  in  deep  water. 


THE  STOMIATIDS.     FAMILIES  GONOSTOMIDAE  AND  STOMIATIDAE 


The  stomiatids  include  many  soft-rayed  fishes 
of  the  mid-depths,  of  most  diverse  appearance,  all 
of  them  with  well  developed  luminescent  organs, 
with  large  eyes,  large  mouths,  and  teeth  in  both 
jaws.  Some  have  and  others  lack  the  adipose 
fin,  but  the  ventrals  are  inserted  more  than 
one- third  of  the  way  back  on  the  abdomen  in  all 
of  them.  They  differ  from  the  herrings  and 
salmons  in  the  structure  of  the  skull.  Four 
species  have  been  taken  in  our  Gulf,  as  strays 
from  offshore. 

Cyclothone  Cyclothone  signata  Garman  1899 

Garman,  Mem.  Mus.  Compar.  Zoology,  vol.  24.  1899, 
p.  246,  pi.  J,  fig.  3. 

Description. — The  general  aspect  of  cyclothone 
is  extremely  characteristic,  the  somewhat  com- 
pressed body  being  deepest  at  the  gill  opening 
with  the  upper  surface  of  the  head  concave  in 
profile,  the  mouth  so  large  that  it  gapes  back  of 
the  eye,  the  lower  jaw  projecting,  the  eye  very 
small,  and  the  gill  openings  very  long.  The 
dorsal  fin  stands  over  the  anal  (the  latter  is  much 

■  Brauer,  Tlefsee-Flsche  Wlssensch.  Ergeb.  Deutschen  Tlefsee-Exped., 
(1898-1899)  1906,  vol.  16,  Pt.  I,  p.  40. 


the  longer  of  the  two),  both  originating  close 
behind  the  middle  of  the  body.  The  caudal 
fin  is  deeply  forked  and  there  is  no  adipose  fin. 

The  luminescent  spots  are  arranged  as  follows: 
One  on  the  head;  1  close  below  the  eye  and  in 
front  of  it;  2  on  each  gill  cover;  9  or  10  between 
the  branchiostegal  rays;  2  longitudinal  rows 
along  each  side  of  the  body,  a  lower  row  of  13 
from  throat  to  ventral  fins,  4  from  ventrals  to 
anal  fin,  and  13  from  anal  to  caudal,  and  an  upper 
row  of  7  reaching  about  as  far  back  as  the  ventrals. 

Color. — Cyclothone  signata  is  colorless  or  pale 
gray,  except  that  the  blackish,  dark  silvery  lining 
of  the  abdominal  cavity  shows  through,  that  the 
luminous  organs  are  black  rimmed  and  silver  cen- 
tered, and  that  there  are  the  following  black 
markings:  a  Y-shaped  mark  on  the  forehead;  a 
series  of  spots  or  short  transverse  stripes  on  the 
flank;  spots  between  the  bases  of  the  dorsal  and 
anal  fin  rays;  one  or  two  transverse  streaks  across 
the  bases  of  the  caudal  fin  rays;  and  a  number  of 


»  Reported  to  us  by  Walter  H.  Rich. 

«  Qoode  and  Bean  (Smithsonian  Contrlb.  Knowl.,  1895,  p.  97)  Ust  these 
captures. 


Figure  64. — Cyclothone  (Cyclothone  aignata) .     After  Brauer. 


FISHES   OF  THE  GULF  OF   MAINE 


147 


irregular  flecks  and  dots  along  the  back  and  on 
the  gill  covers.41 

General  range.- — This  is  an  oceanic  fish,  very 
abundant  in  temperate  latitudes  in  the  Atlantic 
where  it  lives  pelage  from  about  100  fathoms 
down  to  250  fathoms;  hundreds  have  often  been 
taken  in  a  single  haul.  It  is  also  known  from  the 
Pacific. 

Occurrence  in  the  Gulf  of  Maine. — Cyclothone 
appears  within  our  limits  only  as  a  stray  from  the 
Atlantic  Basin;  one  23  mm.  long  that  we  took  in 
a  haul  from  30  fathoms  on  Browns  Bank,  June  24, 
1915,  and  a  second  mutilated  specimen  probably 
of  this  species  from  the  Fundy  Deep  (haul  from 
90  fathoms),  March  22,  1920,  are  the  only  definite 
records. of  it  within  our  limits. 

Stomias  Stomias  ferox  Reinhardt  1842 
Jordan  and  Evermann,  1896-1900,  p.  588. 

Description. — The  members  of  this  genus  (there 
are  several),  resemble  the  viperfish  in  their  bull- 
dog-shaped heads,  with  large  mouth  and  long, 
fanglike  teeth.  But  they  do  not  have  an  adipose 
fin;  the  dorsal  fin  and  the  anal  fin  both  stand  far 
rearward  close  to  the  tail  fin;  the  dorsal  fin  is  even 
with  the  anal  fin;  and  the  first  dorsal  fin  ray  is 
not  prolonged  as  it  is  in  the  viperfish.  The  chin 
bears  a  fleshy  barbel  nearly  as  long  as  the  head 
and  ending  in  a  group  of  about  three  simple  fila- 
ments. The  sides  of  the  body  are  clothed  with 
about  6  rows  of  large,  thin,  somewhat  irregular, 
hexagonal  scales,  and  there  is  one  row  of  lumines- 
cent spots  low  down  along  each  side  and  two  rows 
along  the  belly;  also  one  small,  circular  fight  organ 
below  each  eye. 

The  tip  of  tbe  lower  jaw  overlaps  and  encloses 
the  tip  of  the  upper  jaw  when  the  mouth  is  closed 
in  the  only  member  of  the  genus  that  has  been 
reported  from  the  Gulf  of  Maine  (or  is  likely  to 
be  found  there) ;  the  slender  body  is  about  17  times 


a  For  detailed  accounts  and  colored  Illustrations  see  Garman  (Mem.  Mus. 
Comp.  Zool.,  vol.  24,  1899,  p.  246,  pi.  J,  flg.  3),  Brauer  (Wlssonsch.  Ergeb. 
Deutschen  Tiefsee-Eiped.  (1898-1899),  1908,  vol.  15,  Pt.  1,  p.  77,  pi.  6,  flg.  6), 
Murray  and  HJort  (Depths  of  the  Ocean,  1912,  pi.  I). 


as  long  as  it  is  high;  the  ventral  fins  are  only  about 
as  long  as  the  head;  the  dorsal  fin  is  of  about  the 
same  size  and  shape  as  the  anal  fin,  over  which 
it  stands;  and  there  are  about  85-86  light  organs 
in  each  of  the  ventral  rows,  about  60  fight  organs 
in  each  of  the  lateral  rows. 

Color. — Black  below  as  well  as  above,  the  sides 
with  metallic  iridescence. 

General  range  and  occurrence  in  the  Gulf  of 
Maine. — This  oceanic  fish  is  so  widespread  in  the 
northern  North  Atlantic  that  it  was  taken  at 
almost  all  the  stations  that  the  Michael  Sars 
occupied  there  in  1910,*3  mostly  between  the  75 
to  80  fathom  level  and  the  410  fathom  (750  meter) 
level,  most  plentifully  at  about  275  fathoms  (500 
meters).  The  early  cruises  of  the  Blake  and 
Albatross  I  took  it  at  many  localities  also,  along 
the  continental  slope  of  North  America  between 
the  southeastern  slope  of  the  Newfoundland 
Banks  and  the  Bahama  Channel."  Our  only 
reason  for  mentioning  it  is  that  one  specimen 
about  12  inches  long  (tip  of  snout  to  base  of  tail 
fin)  was  taken  by  a  trawler  on  the  northeastern 
part  of  Georges  Bank  (lat.  42°10'  N.,  long.  67°05' 
W.),  at  about  100  fathoms,  on  January  20,  1936.** 

Stomioides  nicholsi  Parr  1933 

Parr,  Copeia,  1933,  No.  4,  p.  177. 

Description. — The  chief  anatomical  character 
separating  Stomioides  from  Stomias  is  the  struc- 
ture of  the  chin  barbel.  In  Stomias  this  terminates 
in  three  simple  filaments.  But  in  Stomioides  it  not 
only  has  these  barbels,  but  the  main  trunk  ia 
swollen  at  the  tip  and  has  two  additional  filaments 
on  ODe  side  a  little  inward  from  its  tip.  Another 
difference  is  that  the  point  of  origin  of  the  anal  fin 
is  in  advance  of  the  origin  of  the  dorsal  fin  by  a 
distance  about  as  great  as  the  diameter  of  the  eye 
in  Stomioides,  whereas  the  point  of  origin  of  the 
anal  fin  is  about  even  with  that  of  the  dorsal  in 


«  Murray  and  HJort,  Depths  of  the  Ocean,  1912,  pp.  603,  611,  629. 
»  For  a  list  of  these  stations,  see  Goode  and  Bean,  Smithsonian  Contrlb. 
Know!.,  vol.  30, 1898,  p.  107. 
<•  This  specimen  Is  now  In  the  Museum  of  Comparative  Zoology. 


Figure  65. — Stomias  (Stomias  ferox) ,  Banquereau  Bank.     From  Goode  and  Bean.     Drawing  by  H.  L.  Todd. 


148 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


--»<■■>--?"* 


Figoee  66. — Stomioides  nicholsi,  Browns  Bank.     Drawing  by  Myvanwy  M.  Dick. 


Stomias.  Stomioides  resembles  Stomias  in  all  other 
respects  so  closely  that  should  a  specimen  of  either 
be  taken,  that  is  not  easily  identified,  we  suggest 
forwarding  it  to  the  U.  S.  Fish  and  Wildlife  Service 
for  naming. 

Color. — Black  below  as  well  as  above,  the 
luminescent  organs  showing  as  whitish  dots. 

Size. — The  only  specimen  yet  seen  is  about  10% 
inches  long,  from  tip  of  snout  to  base  of  tail  fin. 

Range  and  occurrence  in  the  Gulf  of  Maine. — The 
only  known  specimen  of  this  species  was  taken 
from  the  stomach  of  a  swordfish  harpooned  from 
the  schooner  Barbara,  Capt.  C.  A.  Turner,  on  the 
southeastern  edge  of  Browns  Bank,"  over  the 
250  fathom  line,  August  3,  1932.  Presumably  it 
had  strayed  from  the  mid-depths  offshore. 

Trigonolampa  miriceps  Regan  and  Trewavas  1930 

Regan  and  Trewavas,  Danish  Dana  expeds.  1920-1922, 
No.  6,  1930,  p.  55,  pi.  1,  fig.  1. 

Trigonolampa  resembles  Stomias  in  general 
appearance,  in  the  relative  sizes  and  locations  of 
the  fins,  and  in  having  a  long  fleshy  barbel  on  its 
chin.  But  it  not  only  has  a  small  light  organ  below 
the  eye  (as  in  Stomias),  but  also  has  a  small 
luminescent  patch  close  behind  it,  and  likewise  a 
larger  triangular  patch  extending  from  close  behind 
the  eye  back  across  the  top  of  the  cheek;  these  are 


••This  specimen,  described  by  Parr  (Copela,  1933,  p.  177),  Is  now  In  the 
Museum  of  Comparative  Zoology. 


its  most  distinctive  characters.  The  one  species  of 
the  genus  yet  known  differs  further  both  from 
Stomias  (p.  147)  and  from  Stomioides  (p.  147)  in  a 
considerably  deeper  body  (cf.  fig.  67  with  figs.  65, 
66) ;  also  in  that  the  tip  of  its  lower  jaw  does  not 
enclose  the  tip  of  its  upper  jaw  when  the  mouth  is 
closed;  that  the  point  of  origin  of  its  dorsal  fin  is  in 
advance  of  its  anal  fin  by  a  distance  about  as  great 
as  the  diameter  of  the  eye;  and  that  it  has  only 
about  68  light  organs  in  each  of  its  ventral  rows,  as 
against  85  or  86  in  Stomias  (p.  147). 

Color. — Not  known,  but  probably  black  or  very 
dark  brown.47 

Size.- — The  largest  specimen  yet  seen  (in  the 
Museum  of  Comparative  Zoology)  is  about  9 
inches  (230  mm.)  long  to  the  base  of  the  caudal 
fin. 

Range  and  occurrence  in  the  Gulf  of  Maine. — 
Only  three  specimens  have  been  seen  yet.  The 
first  was  taken  in  the  eastern  Atlantic  by  the 
Danish  research  vessel  Thor  in  1906  at  a  depth  of 
about  600  fathoms;  a  second  was  found  by  Capt. 
John  Toothaker  in  the  stomach  of  a  swordfish 
harpooned  on  the  southern  edge  of  Georges  Bank 
in  the  summer  of  1922,48  and  a  third,  now  in  the 
Museum  of  Comparative  Zoology,  was  recorded 
simply  as  taken  on  Georges  Bank  about  1913.  It 
reaches  the  slope  of  our  outer  Banks  only  as  a 
stray  from  the  mid-depths  offshore. 

"  One  that  we  have  seen  Is  brown  below  as  well  as  above  wherever  the  skin 
Is  intact,  with  the  light  organs  showing  as  darker  dots. 

"  Parr  (Copela,  1933,  No.  4,  p.  178)  has  given  a  detailed  description  of  this 
specimen,  which  is  new  in  the  Museum  of  Comparative  Zoology. 


^■^gf-gl-la;'^'  PI''1 SPPgggjg 


— — ^m^-  ^ 

^^^w 


Figure  67. — Trigonolampa  miriceps.     After  Regan  and  Trewavas. 


FISHES   OF  THE  GULF  OF   MAINE 


149 


HATCHET  FISHES.     FAMILY  STERNOPTYCHIDAE 


These  are  deep,  thin,  flat-sided  little  fishes,  with 
various  spiny  projections,  large  oblique  mouths 
with  small  teeth,  large  eyes  which  are  directed  up- 
wards in  some  of  them  but  sidewise  in  others,  and 
ventral  fins  placed  far  back.  Some  of  them  have 
an  adipose  fin  behind  the  rayed  dorsal,  but  others 
do  not.  All  of  them  are  silvery,  and  all  of  them 
have  series  of  large  and  conspicuous  luminescent 
organs  on  the  lower  part  of  the  body.  They  are 
to  be  found  in  the  mid-depths  in  all  oceans,  some- 
times in  great  abundance. 

Silver  hatchetfish  Argyropelecus  aculeatus 
Cuvier  and  Valenciennes  1849 

Jordan  and  Evermann,  1896-1900,  p.  604,  as  A.  olfersi 
Cuvier,  1829. 


Figure  68. — Silver  hatchetfish  (Argyropelecus  aculeatus). 
After  Brauer. 

Description.- — This  little  fish  is  of  so  bizarre  an 
appearance  that  once  seen  it  could  hardly  be  mis- 
taken for  any  other  species  yet  known  from  our 
Gulf,  or  for  any  that  is  likely  to  stray  thither, 
unless  one  of  its  own  tribe.49  Its  body  is  very 
thin  sidewise,  with  its  forward  part  a  little  less 
than  three-fourths  (70  percent)  as  deep  as  it  is 
long  from  snout  to  base  of  tail  fin,  but  with  the 
ventral  contour  bending  upward  abruptly  about 
midway  of  its  length  in  characteristic  contour,  so 
that  the  rear  half  is  much  less  deep  than  the  for- 
ward half.  This  break  in  the  ventral  contour  is 
marked  by  two  short  bony  spurs,  which  are  out- 

*•  The  several  known  species  of  Argyropelecus  resemble  one  another  so 
closely  that  their  Identification  calls  for  a  specialist.  They  have  been  re- 
viewed by  Schultz,  Proc.  U.  8.  Nat.  Mus.,  vol.  86,  1938,  pp.  147-153.  The 
most  detailed  description  of  this  particular  hatchetfish,  with  the  best  Illus- 
tration (copied  here  as  fig.  68)  Is  by  Brauer,  Wlss.  Ergeb.  Deutschen  Valdicta 
Tlefsee-Exped.,  vol.  18,  Pt.  1, 1908,  p.  110,  fig.  47. 


growths  from  the  pubic  bones,  and  there  is  a  short 
single  spur  (outgrowth  from  the  pectoral  arch)  in 
front  of  them  in  the  mid  ventral  line. 

The  eyes  are  large,  so  high  up  that  the  space 
between  them  on  the  top  of  the  head  is  very 
narrow,  and  they  are  directed  more  upward  than 
sidewise.  The  mouth  is  noticeably  large,  with 
wide  gape,  and  it  is  so  strongly  oblique  that  the 
upper  jaw  is  nearly  vertical.50  The  tips  of  the  two 
jaws  are  about  even  one  with  the  other  when  the 
mouth  is  closed,  and  both  jaws  are  armed  with  a 
large  number  of  tiny  sharp  teeth.  The  dorsal  fin 
is  short,  about  midway  of  the  fish,  and  of  two 
parts,  separated  by  a  deep  but  short  notch.  The 
forward  subdivision  is  in  the  form  of  a  hard, 
triangular  plate  (apex  rearward)  supported  by  8 
or  9  hard  spines,  the  rearmost  of  which  is  the 
stoutest  and  longest.  The  rearward  subdivision 
is  supported  by  9  soft  rays,  that  are  bifid  toward 
their  tips.  The  adipose  fin  is  long  and  low.  The 
pectorals  are  as  long  as  about  two-fifths  the 
greatest  height  of  the  body.  The  ventral  fins, 
each  with  6  soft  rays,  stand  close  behind  the  break 
in  the  ventral  contour  of  the  body,  and  they  are 
connected  with  the  anal  fin  by  a  thin  transparent 
ridge.  The  anal,  commencing  about  under  the 
rear  end  of  the  base  of  the  soft  rayed  part  of  the 
dorsal,  is  notched  midway  of  its  length ;  its  forward 
part  is  supported  by  7  rays  close  together,  the  rear 
part  by  5  shorter  rays  spaced  more  widely.  The 
caudal  fin  is  forked.  A  noticeable  feature  is  that 
the  ventral  edge  of  the  deep  forward  part  of  the 
body,  from  the  pectoral  spur  to  the  pubic  spines, 
is  sharp,  with  a  series  of  12  hard,  plate-like  scales 
or  scutes,  that  extend  for  some  distance  up  the 
sides,  each  slightly  overlapping  the  next  rearward, 
and  the  profile  is  saw-edged  between  the  ventral 
and  anal  fins. 

The  hatchetfishes  are  provided  with  a  complex 
system  of  conspicuous  light-producing  spots.  The 
species  aculeatus  has  one  row  of  12  very  low  down 
along  each  side  of  the  deep  forward  part  of  the  body ; 
also,  a  second  row  higher  up  consisting  of  6  in  front 
of  each  pectoral  fin,  2  along  the  base  of  the  pectoral, 
6  between  pectoral  and  ventral  fins,  4  between  the 
ventral  and  the  anal  fins,  6  along  the  anal,  and  4 
very  small  ones  between  anal  and  tail  fins.  There 
is  also  one  light-organ  a  little  below  and  behind 


1  Most  of  the  published  Illustrations  of  Argyropelecus  fall  to  show  this. 


150 


FISHERY  BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


each  eye,  2  on  the  lower  part  of  the  gill  cover  on 
each  side  and  about  5  on  the  lower  jaw  on  each  side. 

Color. — Perhaps  the  most  conspicuous  feature  of 
the  hatchetfishes,  as  taken  from  the  water,  is  that 
their  entire  bodies  are  glistening  silvery.  On  some 
specimens  the  silver  is  underlaid  with  velvet  black 
over  the  truDk  as  a  whole;  on  others  the  black  un- 
der pigment  is  confined  to  a  marginal  band,  broader 
or  narrower.  The  luminescent  spots  are  pale  yel- 
low or  white. 

Size. — Maximum  length  probably  not  more  than 
3  inches  or  so. 


Range  and  occurrence  in  the  Gulf  oj  Maine. — All 
the  members  of  this  genus  are  oceanic,  and  inhabit 
the  mid-depths.  Localities  listed  by  Schultz  n 
for  this  species  include  the  Grand  Banks,  between 
Georges  and  Browns  Banks,  and  the  offing  of  New 
Jersey  and  Virginia  in  the  western  Atlantic;  the 
Gulf  of  Mexico;  West  Indies;  off  the  South  African 
coast;  the  Indian  Ocean;  and  the  Philippines.  Our 
only  reason  for  mentioning  the  hatchetfish  is  that 
one  specimen  was  taken  on  August  31,  1883,  by  the 
Albatross  between  Georges  and  Browns  Banks 
where  the  depth  was  144  fathoms.62 


THE    EELS.     FAMILIES    ANGUILLIDAE,    CONGRIDAE,   SIMENCHELYIDAE,   SYNAPHO- 
BRANCHIDAE,  NEMICHTHYIDAE,  AND  OPHICHTHYIDAE 


Eels  have  no  ventral  fins;  either  they  have  no 
scales  or  these  are  so  small  as  to  be  hardly  visible; 
their  fins  are  soft,  without  spines ;  the  gill  openings 
are  very  small;  the  vertebrae  extend  in  a  straight 
fine  to  the  tip  of  the  tail ;  and  a  single  fin  runs  over 
the  back,  around  the  tail  and  forward  on  the  belly 
with  no  separation  into  dorsal,  caudal,  and  ventral 
portions.  All  the  species  of  eels  known  from  the 
Gulf  of  Maine  have  pectoral  fins,  but  most  of  the 
morays  of  warmer  seas  are  without  pectorals. 
There  are  several  other  fishes  of  eel-like  form  in  the 
Gulf  of  Maine,  viz.,  the  hag  and  the  lampreys  the 
rock  eel  (Pholis);  the  snake  blenny  (Lumpenus); 
the  wrymouth  (Cryptacanthodes) ;  the  eel  pout 
(Macrozoarces);  and  the  sand  eel  (Ammodytes). 
But  the  jawless,  sucker-like  mouth  of  the  first  two 
separates  them,  at  a  glance,  from  the  true  eels, 
while  there  either  is  a  well-marked  separation  be- 
tween anal  and  caudal  fins  in  all  the  rest;  or  they 
have  ventral  fins  (large  or  small),  or  the  dorsal 
fin  is  spiny,  not  soft. 

Only  five  true  eels  are  known  from  the  Gulf  of 
Maine:  the  common  eel  (p.  151),  the  slime  eel  (p. 


157),  the  conger  (p.  154),  the  snipe  eel  (p.  159),  and 
the  snake  eel  (p.  159),  which  fall  into  five  different 
families  according  to  American  usage.  A  sixth  spe- 
cies, the  long-nosed  eel  (a  deep-water  form  p.  158) 
is  to  be  expected  in  the  deepest  parts  of  the  Gulf 
though  it  has  not  actually  been  recorded  there  as 
yet.  The  group  likewise  includes  the  morays  of 
warm  seas  and  sundry  deep-sea  forms,  some  of 
them  exceedingly  bizarre  in  appearance. 

Common,  conger,  slime,  and  long  nosed  eels  look 
much  alike  in  general  form,  but  are  separated  from 
one  another  by  the  size  of  the  mouth  and  by  the 
relative  lengths  of  the  fins.  In  the  snipe  eels  the 
two  jaws  are  prolonged  into  a  very  long  slender 
beak,  recalling  that  of  a  silver  gar,  the  tail  is  whip- 
like, the  neck  noticeably  slimmer  than  the  head, 
and  the  general  form  extremely  slender,  while  the 
snake  eel  is  very  slender  with  a  hard  pointed  tail. 


•i  Bull.  U.  S.  Nat.  Mus..  vol.  86, 1938,  pp.  146-147  and  pp.  151-152. 

«  Albairos!  station  2063,  lat.  42°  23'  N.,  long.  66°  23'  W.  This  specimen  was 
recorded  by  Qoode  and  Bean  (Smithsonian  Contrlb.  Knowl.,  vol.  30, 1895,  p. 
127)  as  A.  olfersll.  But  Schultz  (Proc.  U.  S.  Nat.  Mus.,  vol.  86, 1938,  p.  151) 
has  found,  on  reexamination,  that  It  Is  an  acuhaiui. 


KEY  TO  GULF  OF  MAINE  EELS 

1.  Both  jaws  are  prolonged  into  a  long  slender  bill Snipe  eel,  p.  159 

The  jaws  are  not  bill-like 2 

2.  The  anal  fin  originates  well  in  front  of  the  point  of  origin  of  the  dorsal  fin Long-nosed  eel,  p.  158 

The  anal  fin  originates  well  behind  the  point  of  origin  of  the  dorsal  fin 3 

3.  The  dorsal  fin  originates  far  behind  the  tips  of  the  pectorals Eel,  p.  151 

The  dorsal  fin  originates  close  behind  the  tips  of  the  pectorals 4 

4.  Mouth  very  small,  its  gape  not  reaching  back  as  far  as  the  eye;  body  very  soft Slime  eel,  p.  157 

Mouth  large,  gaping  back  as  far  as  the  middle  of  the  eye;  body  firm 5 

5.  Mouth  gaping  back  only  about  as  far  as  the  middle  or  rear  edge  of  eye;  body  moderately  stout;  tip  of  tail  soft,  rounded 

Conger,  p.  154 
Mouth  gaping  back  considerably  beyond  eye;  body  very  slender,  tip  of  tail  hard  and  pointed Snake  eel,  p.  159 


FISHES   OF  THE  GULF   OF   MAINE 


151 


Eel  Anguilla  rostrata  (LeSueur)  1817 

American  eel;  Silver  eel;  Fresh-water  eel; 
Elver  (young) 

Jordan  and  Evermann  1896-1900,  p.  348,  A.  ehrysypa 
Rafinesque  1817. 

Description. — In  the  common  American  eel  the 
dorsal  fin  originates  far  behind  the  pectorals, 
this  character  is  enou^j  to  distinguish  it  from  the 
conger,  from  which  it  also  differs  in  that  the  lower 
jaw  projects  beyond  the  tipper  or  at  least  equals 
it  in  length,  and  its  eyes  are  small  and  round. 
Furthermore,  it  develops  scales  as  it  grows,  though 
these  are  so  small  that  they  might  be  overlooked. 
The  eel,  however,  has  a  pointed  snout,  like  the 
conger,  a  large  mouth  gaping  back  as  far  as  the 
middle  of  the  eye  or  past  it;  and  its  gill  slits  are 
set  vertically  on  the  sides  of  the  neck,  their  upper 
corners  abreast  of  the  center  of  the  base  of  the 
pectoral  fin.  It  is  very  closely  related  to  the 
European  eel  (Anguilla  vulgaris),  but  has  fewer 
vertebrae  (average  about  107  as  compared  with 
about  114  or  115  in  the  European  species). 

Color. — The  colors  of  eels  vary  widely  with  the 
bottom  on  which  they  live.  As  a  rule  they  are 
dark  muddy  brown  or  olive-brown  above,  more 
or  less  tinged  with  yellow  on  the  sides;  the  lower 


surface  paler  brown  and  yellower,  with  dirty  yel- 
lowish-white belly.  It  is  common  knowledge 
that  eels  are  dark  if  living  on  dark  mud  but  much 
paler  on  pale  sand.  And  Parker  M  has  found  that 
they  can  change  from  pale  to  dark  in  about  1% 
hours  and  from  dark  to  pale  in  a  little  more  than 
3  hours,  if  moved  from  a  white  background  to  a 
black  or  vice  versa,  under  a  strong  light. 

Size. — Eels  are  said  to  grow  to  4  feet  in  length 
and  to  16K  pounds  in  weight.  Full-grown  females 
average  only  about  2  to  3%  feet,  however,  and 
males  are  smaller.  Any  eel  more  than  18  inches 
long  would  probably  be  a  female,  and  one  more 
than  24  inches  in  length  would  certainly  be  one. 
The  smallest  mature  males  are  about  11  to  12 
inches  long,  females  about  18  inches. 

Habits. — The  life  history  of  the  eel  remained  a 
mystery  until  very  recently.  It  has  been  com- 
mon knowledge  for  centuries  that  young  elvers 
run  up  into  fresh  water  in  spring,  and  adults 
journey  downstream  in  autumn.  A  host  of  myths 
grew  up  to  explain  the  utter  absence  of  ripe  eels 
of  either  sex,  either  in  fresh  water  or  along  the 
seacoast.  But  it  was  only  a  few  years  ago  that 
the  breeding  places  of  the  European  and  American 
eels  were  discovered  and  the  history  of  their  larvae 


«  Jour,  of  Exper.  Zool.,  vol.  88, 1945,  No.  3,  pp.  211-234. 


Figure  69. — Eel  (Anguilla  rostrata).  A,  adult,  Connecticut  River,  Massachusetts;  from  Goode,  drawing  by  H.  L. 
Todd;  B,  "Leptocephalus"  stage,  49  mm.;  C,  "Leptocephalus"  stage,  55  mm.;  D,  "Leptocephalus"  stage,  58  mm.; 
E,  transformation  stage,  61  mm.  B-E,  after  Schmidt. 


152 


FISHERY   BULLETIN   OF   THE    FISH   AND   "WILDLIFE   SERVICE 


traced,  chiefly  by  the  persevering  researches  of 
the  Danish  scientist,  Johannes  Schmidt.64  Now 
we  know  that  the  life  history  of  the  eel  is  just  the 
antithesis  of  that  of  the  salmon,  shad,  and  alewife, 
for  eels  breed  far  out  at  sea,  but  make  their  growth 
either  in  estuarine  situations  or  in  fresh  water. 

The  young  elvers,  averaging  from  2  to  3%  inches 
in  length,  appear  along  our  shores  in  spring.  As 
yet  we  have  few  data  on  the  exact  date  of  their 
arrival  on  the  Gulf  of  Maine  coast.  They  appear 
as  early  as  March  at  Woods  Hole;  by  mid-  or 
late  April  both  in  Narragansett  Bay  and  in  Passa- 
maquoddy  Bay  at  the  mouth  of  the  Bay  of  Fundy, 
while  Welsh  encountered  a  tremendous  run  in 
Little  River,  near  Gloucester,  on  May  5,  1913, 
suggesting  that  they  may  be  expected  in  the 
mouths  of  most  Gulf  of  Maine  streams  during 
that  month.  And  they  are  found  ascending 
streams  in  the  Bay  of  Fundy  region  during  the 
summer.  A  run  may  last  for  a  month  or  more  in 
one  stream,  only  for  a  few  days  in  another.  And 
there  is  a  noticeable  segregation  even  at  this  early 
stage,  some  of  the  elvers  remaining  in  tidal 
marshes,  in  harbors,  in  bays  back  of  barrier 
beaches,  and  in  other  similar  situations,  some  even 
along  the  open  coast,  especially  where  there  are 
beds  of  eel  grass  (Zostera);  while  others  go  into 
fresh  water,  some  of  them  ascending  the  larger 
rivers  for  tremendous  distances.65 

It  is  now  generally  believed  that  most  of  the 
eels  that  are  caught  in  fresh  water  are  females. 
But  some  of  the  females  remain  in  salt  marshes 
and  harbors,  to  judge  from  the  large  size  of  many 
of  the  eels  that  are  caught  there.  And  nothing  is 
known  as  to  what  preference  the  males  of  the 
American  eel  may  show  in  this  respect. 

It  is  no  wonder  that  the  ability  of  the  elvers  to 
surmount  obstacles  as  they  run  upstream  is 
proverbial,  for  they  clamber  over  falls,  dams,  and 
other  obstructions,  even  working  their  way  up 
over  damp  rocks  as  Welsh  saw  them  doing  in 
Little  River,  where  they  were  so  plentiful  on  May 
5  and  7,  1913,  that  he  caught  1,500  in  one  scoop 


«  The  life  history  of  the  eel  Is  presented  In  more  detail  than  Is  possible  here 
by  Schmidt  (PhUos.  Trans.  Roy.  Soc.  London,  Series  B,  vol.  211  (1922) 
1923,  pp.  179-208,  summarized  In  Nature,  vol.  110, 1922,  p.  716),  and  by  Cun- 
ningham (Nature,  vol.  113, 1924,  p.  199).  See  also  Schmidt  (Rapp.  et  Proc- 
Verb.  Cons.  Perm.  Internat.  Explor.  Mer,  vol.  5,  No.  4,  1906,  pp.  137-204, 
pis.  7-13);  for  a  popular  account  see  Smith  (Nat.  Qeog.  Mag.,  vol.  24,  No.  10, 
October  1913,  p.  1140). 

u  Eels  are  native  In  Lake  Ontario  which  they  reach  by  way  of  the  St. 
Lawrence  River;  and  up  the  Mississippi  drainage  systems  even  as  far  as 
North  Dakota,  Wisconsin,  Ohio,  and  western  Pennsylvania. 


of  a  small  dip  net  and  545  with  a  few  grabs  of 
his  bare  hand.  Elvers  in  equal  multitudes  have 
often  been  described  in  other  streams,  American 
as  well  as  European.  Eels  can  live  out  of  water 
so  long  as  to  give  rise  to  the  story  that  they  often 
travel  overland.  There  is  no  positive  evidence 
for  this.  But  Sella  6S  has  proved,  by  experiments 
with  European  eels  marked  so  as  to  be  recognizable 
if  recaught,  that  they  can  carry  out  journeys  as 
long  as  31  miles  (50  kilometers)  along  underground 
waterways.  Doubtless  it  is  this  ability  that 
explains  the  presence  of  eels  in  certain  ponds  that 
have  no  visible  outlet  nor  inlet,  a  fact  often 
attested. 

It  is  true  in  a  general  way  that  eels  seek  muddy 
bottom  and  still  water,  as  has  been  said  so  com- 
monly. But  this  is  not  always  so  whether  in  salt 
water  or  in  fresh.  Thus  the  rocky  pool  at  the 
outer  end  of  the  outlet  from  Little  Harbor, 
Cohasset,  on  the  south  shore  of  Massachusetts 
Bay,  is  a  good  place  to  catch  eels;  and  large  ones 
are  only  too  common  in  swiftflowing,  sandy  trout 
streams  on  Cape  Cod ;  we  have  had  one  follow  and 
nibble  at  the  trout  we  were  dragging  behind  us 
on  a  line.  The  fact  is,  they  can  live  and  thrive 
wherever  food  is  to  be  had,  which  applies  to  them 
in  estuarine  situations  and  in  fresh  water. 

No  animal  food,  living  or  dead  is  refused,  and 
the  diet  of  the  eels  in  any  locality  depends  less  on 
choice  than  on  what  is  available.  Small  fish  of 
many  varieties,  shrimps,  crabs,  lobsters,  and 
smaller  Crustacea,  together  with  refuse  of  any 
kind  (for  they  are  scavengers)  make  up  the  bulk 
of  the  diet  in  salt,  estuarine,  and  brackish  water. 
Being  very  greedy,  any  bait  will  do  to  catch  an 
eel.  They  are  chiefly  nocturnal  in  habit,  as  every 
fisherman  knows,  usually  lying  buried  in  the  mud 
by  day  to  venture  abroad  by  night.  But  eels, 
large  and  small,  are  so  often  seen  swimming  about, 
and  so  often  bite  by  day  that  this  cannot  be  laid 
down  as  a  general  rule. 

Eels  tolerate  a  wide  range  of  temperature.  But 
it  is  common  knowledge  that  those  inhabiting  the 
salt  marshes  and  estuaries  of  our  Gulf,  and  its 
tributary  streams,  mostly  lie  inactive  in  the  mud 
during  the  winter. 

Eels  grow  slowly.  Hildebrand  and  Schroeder  67 
concluded  from  a  series  of  measurements  taken  at 
different  seasons  in  lower  Chesapeake  Bay  that  those 

*  Mem.  R.  Comlt.  Talassogr.  Ital.,  vol.  158, 1929. 
»'  Bull.  U.  S.  Bur.  Fish.,  vol.  43, 1928,  p.  114. 


FISHES   OF  THE  GULF  OF   MAINE 


153 


2}j  inches  long  in  April  are  about  5  inches  long  a 
year  later,  or  about  2  years  after  their  transforma- 
tion. The  winter  rings  on  the  scales  have  shown 
that  full  grown  adults  of  the  European  species  are 
from  5  to  20  years  old,  depending  on  food  supply, 
and  other  conditions;  this  is  corroborated  for  the 
American  species  by  the  fact  that  Dr.  Hugh  M. 
Smith,  former  Commissioner  of  the  United  States 
Bureau  of  Fisheries,  found  that  a  female,  on  the 
way  down  the  Potomac,  was  in  her  twelfth  year. 

At  the  approach  of  sexual  maturity,  which  takes 
place  in  the  fall,  the  eels  that  are  in  fresh  water 
drop  downstream,  traveling  mostly  at  night. 
They  now  cease  feeding,  as  do  those  that  have 
been  living  in  the  river  mouths,  bays,  and  estuaries; 
the  color  of  the  back  changes  from  olive  to  almost 
black,  the  ventral  side  turns  silvery,  and  the  eyes 
of  the  males  grow  to  twice  their  previous  size. 
Both  males  and  females  then  move  out  to  sea, 
and  it  is  not  until  after  they  reach  salt  water  that 
the  ovaries  mature.  In  fact,  no  perfectly  ripe 
female  eel  has  ever  been  seen,  and  only  one  ripe 
male  (of  the  European  species). 

So  little  is  the  life  history  of  the  eel  understood 
by  our  fishermen,  that  we  again  emphasize  the 
undoubted  fact  that  no  eel  ever  spawns  in  fresh 
water. 

The  eels  drop  wholly  out  of  sight  when  once 
they  leave  the  shore;68  no  one  knows  how  deep 
they  swim,  but  they  certainly  journey  out  beyond 
the  continental  slope  into  the  oceanic  basin  before 
depositing  their  eggs.  Schmidt  has  been  able  to 
outline  the  chief  spawning  center  of  the  American 
species  (from  the  captures  of  its  youngest  larvae) 
as  between  latitudes  20°  and  30°  N.  and  between 
longitudes  60°  and  78°  W.;  i.  e.,  east  of  Florida 
and  of  the  Bahamas  south  of  Bermuda.  But  it  may 
also  spawn  (always  in  deep  water)  farther  north 
as  well." 

The  American  eel  spawns  in  midwinter,  thus 
occupying  one  to  two  months  in  its  journey  from 
the  coast  to  the  spawning  ground,  for  Schmidt 
found  very  young  larvae  (7  to  8  mm.)  in  February. 
Eels,  like  Pacific  salmon,  die  after  spawning,  the 
evidence  of  this  beiag  that  no  spent  eels  ha  ye  ever 
been  seen  and  that  large  eels  have  never  been 

IS  Large  eels,  on  their  seaward  journey,  have  occasionally  been  caught  by 
otter  trawlers  in  the  western  part  of  the  British  Channel,  but  we  know  of  no 
such  occurrence  on  this  side  of  the  Atlantic. 

«  See  Schmidt  (Ann.  Rep.  Smithsonian  Inst.,  (1924)  1925,  pp.  279-314)  for 
a  readable  account  of  the  investigations  which  enabled  him  to  chart  the 
breeding  places  and  seasons  of  the  American  and  European  eels. 
210941—53 11 


known  to  run  upstream  again.  Smith  suggests 
that  they  probably  "jellify"  and  disintegrate, 
as  the  conger  does. 

Eels  (European)  are  among  the  most  prolific 
fish,  ordinary  females  averaging  5  to  10  million 
eggs  and  the  largest  ones  certainly  15  to  20 
million.  It  is  doubtful  whether  eggs  laid  by  the 
American  eel  have  been  seen,  or  of  the  European 
either,  for  that  matter.60  But  it  is  generally 
supposed  that  they  float  in  the  upper  or  inter- 
mediate water  layers  until  hatching.  The  larval, 
so-called  "leptocephalus"  stage,  like  that  of  all 
the  true  eels,  is  very  different  in  appearance  from 
the  adult,  being  ribbon-like  and  perfectly  trans- 
parent, with  small  pointed  head;  and  it  has  very 
large  teeth,  though  it  is  generally  believed  to  take 
no  food  until  the  time  of  its  metamorphosis. 
These  leptocephali  of  our  eel,  living  near  the 
surface,  have  been  found  off  our  coasts  as  far  north' 
as  the  Grand  Banks,  but  never  east  of  longitude 
50°  W. 

Inasmuch  as  the  breeding  areas  of  the  American 
and  European  eels  overlap,  not  the  least  inter- 
esting phase  of  the  lives  of  the  two  is  that  the 
larvae  of  the  American  species  should  work  so 
consistently  to  the  western  side  of  the  Atlantic, 
and  those  of  the  European  to  the  eastern  side 
that  no  specimen  of  the  former  has  ever  been 
taken  in  Europe  or  of  the  latter  in  America. 

The  American  eel  takes  only  about  one-third 
as  long  as  the  European  to  pass  through  its  larval 
stage;  i.  e.,  hardly  a  year,  as  against  2  to  3  years. 
The  leptocephali  reach  their  full  length  of  60  to 
65  mm.  by  December  or  January,  when  meta- 
morphosis takes  place  to  the  "elver";  the  most 
obvious  changes  being  a  shrinkage  in  the  depth 
and  length  of  the  body  but  an  increase  in  its 
thickness  to  cylindrical  form,  loss  of  the  larval 
teeth,  and  total  alteration  in  the  aspect  of  head 
and  jaws,  while  the  digestive  tract  becomes 
functional. 

It  is  not  until  they  approach  our  shores,  how- 
ever, that  the  adult  pigmentation  develops  or 
that  the  elver  begins  to  feed,  a  change  that  is 
accompanied  by  a  second  decrease  in  size.  How 
such  feeble  swimmers   as   the  leptocephali  find 

"  Four  eggs  taken  on  the  Arcturus  expedition  near  Bermuda  in  1925  wer» 
provisionally  identified  as  those  of  the  American  eel  by  Fish  who  has  pictured 
them  and  the  larvae  hatched  from  one  of  them  (Zoologica,  New  York  Zool. 
Soc,  vol.  28,  1927,  pp.  290-293,  flgs.  103-107).  But  the  date  at  which  they 
were  taken  (July  15-17)  makes  it  more  likely  that  they  belonged  to  some  other 
member  of  the  eel  tribe. 


154 


FISHERY   BULLETIN    OF   THE    FISH    AND    WILDLIFE    SERVICE 


their  way  into  the  neighborhood  of  the  land 
remains  a  mystery.  It  seems  certain,  however, 
that  all  the  young  eels  bound  for  the  Gulf  of 
Maine  complete  the  major  part  of  their  meta- 
morphosis while  they  still  are  far  offshore.  Thus 
we  have  never  taken  one  in  the  leptocephalus 
stage  in  the  Gulf  of  Maine  in  all  our  tow-nettings, 
whereas  (more  significant)  the  Albatross  towed 
three  young  eels  in  the  so-called  "glass-eel"  stage, 
54  to  59  mm.  long,  of  practically  adult  form  but 
still  transparent,  during  her  spring  cruise  in  1920, 
one  of  them  on  Georges  Bank,  March  1 1 ;  a  second 
on  Browns  Bank,  April  16;  and  one  in  the  western 
basin  of  the  Gulf  off  Cape  Ann,  February  23. 
Evidently  they  were  intercepted  on  their  way  in 
to  land.  And  since  all  three  were  on  the  surface, 
we  may  take  it  that  glass  eels,  like  leptocephali, 
keep  to  the  uppermost  water  layers  during  their 
journey. 

General  range. — Coasts  and  streams  of  West 
Greenland,81  eastern  Newfoundland,62  Strait  of 
Belle  Isle,  and  northern  side  of  the  Gulf  of  St. 
Lawrence  south  to  the  Gulf  of  Mexico,  Panama, 
West  Indies,  and  (rarely)  to  the  northern  coast 
of  South  America;  also  Bermuda;  running  up  into 
fresh  water  but  going  out  to  sea  to  spawn  p.  153. 

Occurrence  in  the  Gulf  of  Maine. — The  occur- 
rence of  the  eel  around  the  periphery  of  our  Gulf 
can  be  described  in  the  one  word  "universal." 
There  is,  we  believe,  no  harbor,  stream  mouth, 
muddy  estuary,  or  tidal  marsh  from  Cape  Sable 
on  the  east  to  the  elbow  of  Cape  Cod  on  the  west 
but  supports  eels  in  some  numbers,  and  they  run 
up  every  Gulf  of  Maine  stream,  large  or  small, 
from  which  they  eventually  find  their  way  into 
the  ponds  at  the  headwaters  unless  barred  by 
insurmountable  barriers  such  as  very  high  falls. 
Examples  of  long  journeys  by  eels  upstream,  in 
New  England  rivers,  are  to  the  Connecticut  Lakes, 
New  Hampshire,  at  the  head  of  the  Connecticut 
River;  to  the  Rangeley  Lakes  at  the  head  of  the 
Androscoggin,  and  to  Matagamon  Lake,  at  the 
head  of  the  East  Branch  of  the  Penobscot.  Eels 
are  even  caught  in  certain  ponds  without  outlets, 
as  noted  above  (p.  152).  On  the  other  hand,  we 
have  seen  a  few  (and  some  large  ones)  along  the 
open  coast,  at  Cohasset,  for  example,  but  always 

•>  Jensen  (Invest,  of  the  Dana  In  West  Greenland  Waters,  1926,  Eitr. 
Rapp.  et  Proc.  -Verb  Cons.  Internet.  Eipl.  Mer,  vol.  39, 1926,  p.  101)  records 
the  American  eel  as  one  of  the  four  fresh-water  fishes  known  from  the  west 
coast  of  Greenland. 

u  Reported  by  Dr.  O.  W.  Jeffers  as  common. 


close  in  to  the  shore  line  and  in  only  a  few  feet  of 
water,  where  flounder  fishermen  catch  them  from 
time  to  time. 

Importance. — Schmidt  has  suggested  that  the 
American  eel  is  not  as  plentiful  in  actual  numbers 
as  the  European,  arguing  from  the  facts  that  its 
larvae  have  not  proven  so  common  on  the  high 
seas,  and  that  the  American  catch  of  eels  (about 
2,000  tons  yearly)  was  but  a  fraction  as  large  as  the 
European  catch  (about  10,000  tons  annually). 
But  it  is  not  safe  to  draw  any  conclusions  from  the 
statistics  because  the  American  catch  is  limited 
more  by  the  fact  that  eels  are  not  much  in  demand, 
than  by  the  available  supply.  And  the  local 
demand  is  less  for  them  today  than  it  was  30 
years  ago,  as  is  reflected  in  a  decrease  in  the  re- 
ported landings  from  about  305,000  pounds  for 
Maine  and  about  240,000  pounds  for  Massachu- 
setts in  1919  to  about  19,000  pounds  for  Maine 
and  about  32,000  pounds  for  Massachusetts  in 
1947.  The  yearly  landings  of  eels  along  the  Cana- 
dian shore  of  our  Gulf  and  from  the  tributary 
fresh  waters  are  30,000^40,000  pounds  nowadays. 

Practically  the  entire  coastwise  catch  is  made 
in  salt  marshes,  estuaries  and  stream  mouths;  the 
numbers  captured  up  stream  are  negligible  of 
recent  years,  except  in  New  Brunswick  where 
16,000  pounds  were  caught  in  the  lower  sections  of 
the  St.  John  River  System  in  1950.63  In  Germany, 
however,  where  the  demand  for  eels  is  much  greater, 
the  yearly  catch  is  nearly  four  times  as  great  for 
rivers  and  other  fresh  waters  as  it  is  for  the  coast. 
And  many  millions  of  elvers  were  transplanted, 
during  the  1930's,  from  British  rivers  (the  Severn 
in  particular)  to  landlocked  bodies  of  water  in 
Central  Europe  which  the  young  eels  could  not 
reach  naturally. 

The  greater  part  of  the  catch  is  made  in  nets 
and  eelpots;  and  some  are  speared,  mostly  in  late 
autumn  and  winter,  often  through  the  ice. 

American    conger    Conger    oceanica     (Mitchill) 
1818  M 

Sea  eel 

Jordan  and  Evermann,  Leptocephalus  conger  (Linnaeus 
1758),  189&-1900,  p.  354. 

•*  Information  from  A.  H.  Lelm. 

•*  The  American  conger  had  long  been  considered  Identical  with  the  Euro- 
pean. But  Schmidt  (Nature,  vol.  128,  1931,  p.  602)  has  recently  shown 
that  it  is  a  distinct  species,  characterized  by  having  fewer  vertebrae;  a  rela- 
tionship paralleling  that  between  the  American  and  European  eels  of  the 
genus  Anouilla. 


FISHES   OF  THE  GULF  OF   MAINE 


155 


Figure  70. — American  conger  (Conger  oceanica).     A,  adult,  Connecticut;  from  Goode,    drawing   by   H.    L.  Todd;  B, 

"Leptocephalus"  stage,  84  mm.,  Chesapeake  Bay. 


Description. — The  readiest  characters  by  which 
to  distinguish  the  conger  from  other  eels  are  noted 
in  the  key  (p.  150);  notably  the  origin  of  the  dorsal 
fin  above  or  only  very  slightly  behind  the  tip  of  the 
pectoral  when  the  latter  is  laid  back,  the  rather 
long-pointed  snout,  the  large  mouth  cleft  back  at 
least  as  far  as  the  middle  of  the  eye,  and  the 
scaleless  skin.  The  conger  has  many  more  verte- 
brae than  the  common  eel  and  there  are  other 
skeletal  differences.66  The  conformation  of  the  tip 
of  the  snout  likewise  helps  to  identify  the  conger, 
for  its  upper  jaw  usually  projects  beyond  the 
lower,  whereas  in  the  common  eel  the  reverse  is 
true,  or  at  least  the  lower  equals  the  upper.  Fur- 
thermore, the  eyes  of  the  conger  are  oval  anil 
larger  than  the  round  eyes  of  the  common  eel. 

To  give  an  idea  of  the  proportions  of  the  conger, 
we  need  only  add  that  the  distance  from  tip  of 
snout  to  dorsal  fin  is  about  one-fifth  of  the  total 
length;  the  length  of  the  snout  is  one-fourth  that 
of  the  head;  the  length  of  the  pectorals  is  equal  to 
one-third  to  one-fourth  of  the  distance  from 
dorsal  fin  to  tip  of  snout;  and  that  the  body  is  of 
the  snake-like  form  characteristic  of  eels  in  general. 

Color. — Bluish  gray  or  grayish  brown  above, 
sometimes  of  a  reddish  tinge,  sometimes  almost 
black;   paler  on   the   sides;   dingy   white   below. 

Size.  —  This  is  a  much  larger  fish  than  the  com- 
mon  eel.     The   larger   ones   taken   off   southern 


New  England  and  New  Jersey  are  said  to  measure 
4  feet  up  to  7  feet  in  length.  The  general  run  of 
those  caught  weigh  4  to  12  pounds,  the  heaviest 
we  have  seen  weighed  about  22  pounds.  But 
the  North  American  species  never  attains  the 
enormous  size  reached  by  the  European  species;  the 
largest  European  conger  reliably  reported,  of 
which  we  have  read,  was  9  feet  long,  and  weighed 
160  pounds.69 

Habits. — The  depth  range  of  the  conger  is  from 
close  to  the  coastline  (they  are  caught  from  the 
dock  at  Woods  Hole)  out  to  the  edge  of  the 
continental  shelf,  the  deepest  record  for  it  being 
for  one  that  we  trawled  at  142  fathoms  off  southern 
New  England,  on  the  Albatross  III,  in  May  1950. 
It  feeds  chiefly  on  fish:  butterfish,  herring,  and 
eels  have  been  found  in  their  stomachs  at  Woods 
Hole.  They  also  prey  on  shrimps  and  small 
mollusks  at  times.  And  we  have  caught  them 
(and  have  seen  them  caught)  on  crabs,  on  soft 
clams  (Mya),  on  sea  clams  (Mactra)  and  on  cut 
fish  bait. 

It  is  now  well  established  that  the  European 
species  (hence  no  doubt  the  American  also) 
breeds  but  once  during  its  life  and  then  perishes 
like  the  common  eel.  Ripe  congers  are  never 
caught  on  hook  and  line,  for  they  cease  to  feed, 
hence  to  bite,  for  some  time  previous.  But  the 
males  of  the  European  species,  kept  in  aquaria, 


••  For  an  account  of  these,  see  Smltt  (Scandinavian  Fishes,  vol.  2,  1898 
pp.  1016-1017, 1037. 


■  Jenkins,  Fishes  of  the  British  Isles,  1925,  p.  275;  see  also  Day,  Fishes  of 
Great  Britain,  vol.  2  ,1884,  p.  253,  for  large  European  congers. 


156 


FISHERY   BULLETIN   OP   THE    FISH   AND   "WILDLIFE   SERVICE 


have  repeatedly  been  known  to  become  fully  ripe, 
females  nearly  so,67  then  invariably  dying.  The 
ripening  of  the  sexual  products  is  accompanied 
by  changes  in  the  shape  of  the  head ;  in  the  loss  of 
the  teeth;  and  in  a  jellification  of  the  bones,  while 
the  eyes  of  the  males  become  enormous  and  the 
females  become  much  distended  by  the  ovaries. 
It  is  probable  that  the  American  conger  ripens 
off  the  coast  of  southern  New  England  in  summer; 
European  congers  in  captivity  have  been  known 
to  do  so  every  month  in  the  year  except  October 
and  November. 

It  seems  that  the  conger,  like  the  common  eel, 
moves  out  from  the  coast  to  spawn,  for  its  young 
larvae  have  never  been  taken  inshore,  and  Dr. 
Johannes  Schmidt's 68  discovery  of  very  young 
larvae  in  the  West  Indian  region,  but  nowhere 
else,  points  to  this  as  the  chief  spawning  ground 
of  the  American  conger,  if  not  the  only  one. 

The  congers  are  extremely  prolific  fish,  the 
number  of  eggs  a  European  female  may  produce 
having  been  estimated  as  high  as  3  to  6  millions. 
American  conger  eggs  have  never  been  identified, 
for  although  eggs  taken  over  the  tilefish  grounds 
30  miles  south  of  Nantucket  lightship  in  July 
1900  69  have  been  credited  to  this  species,  there  is 
no  certainty  that  this  was  their  true  parentage. 

It  has  long  been  known  that  the  congers,  like 
the  common  eels,  pass  through  a  peculiar  ribbon- 
like  larval  stage  (the  so-called  "leptocephalus") 
very  broad  and  thin  and  perfectly  transparent, 
with  a  very  small  head.70  In  fact  the  first  lepto- 
cephalus ever  seen  (about  1763)  was  the  larval 
European  conger.  But  its  identity  was  not  estab- 
lished definitely  until  1886,  when  the  famous 
French  zoologist,  Delage,71  reared  one  through 
its  metamorphosis  at  the  biological  station  at 
Roscoff. 

The  leptocephalus  stage  of  the  conger  is  rela- 
tively more  slender  than  that  of  the  common 
eel,  it  grows  larger  (to  a  length  of  150-160  mm.), 
and  its  vertebrae  and  muscle  segments  are  far 
more  numerous  (140-149  in  the  American  conger, 

*  Cunningham  (Jour.  Mar.  Biol.  Assoc.  United  Kingdom,  N.  Ser.,  vol.  2, 
1891-92,  pp.  16-12)  gives  an  Interesting  account  of  this  and  other  phases  of  the 
life  history  of  the  conger. 

»  See  Nature,  vol.  128, 1931,  p.  602,  for  a  discussion  of  this  question  by  Dr. 
Schmidt. 

«  Eigenmann,  Bull.  U.  S.  Bur.  Fish.,  vol.  21, 1902,  p.  37. 

'•  For  photographs  of  the  leptocephalus  stage  of  the  European  conger,  see 
Schmidt,  Rapp.  et  Proc.  Verb.  Cons.  Perm.  Intemat.  Eiplor.  Mer,  vol.  5, 
No.  4, 1906,  pi.  9,  flgs.  8,  9;  and  Meddelelser  Komm.  Havundersiigelser,  Ser. 
Flskerl,  vol.  3,  No.  6,  pi.  1,  flgs.  1-3. 

"  Conptes  Rendus  Acad.  Scl.  Paris,  vol.  103, 1886,  p.  698. 


154-163  in  the  European)  than  in  the  common 
eels  (about  107  in  the  American  eel  and  about 
114  in  the  European).  But  the  number  of  body 
segments  (visible  only  under  a  lens)  is  not  of  itself 
a  safe  clue  to  identity,  for  there  are  as  many  or 
more  in  the  long-nosed  eel  (p.  158)  which  has  been 
reported  in  the  Gulf;  also  in  the  mo  rays,  and  in 
various  other  members  of  the  eel  tribe.72 

The  duration  of  the  larval  period  of  the  conger 
is  not  known.  The  process  of  metamorphosis 
consists  essentially  in  a  thickening  and  narrowing 
of  the  body,  an  enlargement  of  the  head,  the  for- 
mation of  the  swim  bladder  and  permanent  teeth, 
and  the  development  of  pigment  in  the  skin,  a 
change  that  occupied  about  two  months  (May  to 
July)  in  the  case  of  Delage's  European  specimen. 
His  young  conger  was  9.3  centimeters  (3.6  inches) 
long  at  its  completion.73 

General  range. — Continental  shelf  of  eastern 
America:  adults  are  known  north  to  the  tip  of 
Cape  Cod;  larval  stages  to  eastern  Maine.  Its 
southern  boundary  cannot  be  stated  until  the 
congers  of  the  coasts  of  North  and  of  South 
America  have  been  critically  compared.  It  is  rep- 
resented by  a  closely  allied  species  (Conger  conger) 
in  the  eastern  North  Atlantic. 

Occurrence  in  the  Gulf  of  Maine. — The  only 
records  for  adult  congers  within  the  limits  set  here 
for  the  Gulf  of  Maine  are  of  one  taken  at  North 
Truro,  Cape  Cod;  a  second  specimen  trawled 
close  to  Provincetown  in  Cape  Cod  Bay,  July  5, 
1951,  by  Capt.  Herman  Tasha;  and  a  third, 
trawled  south  of  Nantucket  shoals  by  Albatross 
III,  in  mid  May  1950.7*  But  the  conger  must  be 
much  more  plentiful  at  times  off  the  shoals  than 
the  foregoing  would  suggest  for  Capt.  Henry 
Klim  of  the  dragger  Eugene  H  reports  trawling 
1 ,400  pounds  of  them  there,  at  76  fathoms,  March 
25-30,  1951."  And  its  curious  band-like  "lepto- 
cephalus" larvae  have  been  found  within  the  Gulf 
on  several  occasions.  Thus,  half  a  dozen  speci- 
mens were  picked  up  on  the  beach  at  Cherryfield 
and  Old  Orchard,  Maine,  and  at  Nahant,  Mass., 


»  Fish  (Zoologlca,  New  York  Zool.  Soc,  vol.  8,  1927,  pp.  307-308)  gives  a 
table  of  the  numbers  of  body  segmentsfor  various  eels  and  for  "leptocephalus" 
larvae  of  known  and  unknown  parentage. 

»  Schmidtleln  (Mittlell,  Zool.  Stat.  Neapel,  vol.  I,  1879,  p.  135)  speaks  of 
young  "congers"  at  Naples  In  April  as  hardly  one-third  as  long  as  this,  a 
discrepancy  suggesting  that  these  may  actually  have  belonged  to  one  of  the 
Muraenold  eels. 

»  Local  reports  of  congers  do  not  necessarily  relate  to  the  true  conger,  for  the 
eel  pout  (p.  510),  which  Is  common  In  the  Gulf,  Is  often  misnamed  thus. 

"  At  lat.  40°  N.,  long.  69°  50'  W. 


FISHES   OF  THE  GULF  OF   MAINE 


157 


more  than  a  half  century  ago.  Two  specimens, 
also  picked  up  on  the  beach,  were  sent  up  from 
Newburyport,  Mass.,  in  November  1929;  and 
A.  H.  Clark,  of  the  U.  S.  National  Museum, 
informs  us  that  he  has  found  many  larvae  of  the 
leptocephalus  type  at  Manchester,  Mass.,  which 
probably  were  congers  to  judge  from  their  size. 

The  conger  occurs  regularly  and  commonly  to 
the  west  and  south  of  Cape  Cod,  being  taken  near 
Woods  Hole  from  July  into  the  autumn,  and  about 
Block  Island  from  August  until  November.  Very 
little  is  known  about  their  movements.  But  we 
suspect  that  they  shift  offshore  into  deeper  and 
warmer  water  for  the  winter,  judging  from  their 
absence  then  in  shoal  water,  contrasted  with  the 
large  offshore  catch  in  March  mentioned  above 
(p.  156)  and  with  the  fact  that  we  saw  several 
trawled  at  50  to  142  fathoms  off  southern  New 
England  on  the  Albatross  III,  in  May  in  1950. 

Slime  eel  Simenchelys  parasiticus  Gill  1879 

Snub-nosed  eel 

Jordan  and  Evermann,  1896-1900,  p.  349. 

Description.- — The  most  distinctive  characters 
of  the  slime  eel,  its  eel-like  form,  snub  nose,  long 
dorsal  fin,  and  soft  and  slimy  body,  have  been 
mentioned  already  (p.  150).  It  is  stouter  and  more 
sway-bellied  than  the  common  eel,  very  soft,  and 
with  a  more  tapering  tail.  The  dorsal  fin  origi- 
nates a  short  distance  behind  the  tips  of  the 
pectorals  when  the  latter  are  laid  back  against  the 
body,  and  the  anal  runs  forward  on  the  lower 
surface  almost  to  the  vent,  which  is  situated  about 
midway  of  the  body.  The  head  is  much  shorter 
than  in  either  the  common  eel  or  the  conger;  the 
mouth  is  small,  gaping  back  only  about  half 
way  to  the  forward  edge  of  the  eye,  with  upper  and 


lower  jaws  of  equal  length  and  each  armed  with  a 
single  series  of  small,  close-set  cutting  teeth.  The 
gill  openings  are  small,  and  instead  of  being 
vertical  and  on  the  sides  of  the  neck  as  they  are  in 
the  common  eel,  they  are  longitudinal  and  lower 
down  on  the  throat. 

Color. — Dark  brown,  with  the  belly  only  a  little 
paler  than  the  back,  though  usually  more  or  less 
silvery. 

Size. — About  2  feet  long. 

Habits. — It  is  partly  parasitic  in  habit,  burrow- 
ing into  the  bodies  of  halibut  and  other  large  fish, 
circumstances  under  which  a  considerable  number 
of  specimens  have  been  brought  in  by  fishermen. 
Very  likely  it  was  common  inshore  in  the  old  days 
when  halibut  were  plentiful  there.  It  also  lives 
independently  on  the  bottom.  Nothing  is  known 
of  its  manner  of  life  beyond  this,  nor  of  its  breeding 
habits.  We  may  add  from  experience  that  it  is  as 
slimy  as  a  hag  and  drips  with  sheets  of  mucus  when 
drawn  out  of  the  water. 

General  range. — The  continental  slope,  and  the 
slopes  of  the  offshore  banks,  from  abreast  of  the 
eastern  end  of  Long  Island  to  the  Newfoundland 
Banks,  in  depths  ranging  from  200  to  more  than 
900  fathoms;  also  in  deep  water  about  the  Azores, 
and  represented  in  Japanese  waters  by  an  ex- 
tremely close  relative,  if,  indeed,  it  is  separable  at 
all  from  the  Atlantic  slime  eel.76 

Occurrence  in  the  Gulf  of  Maine. — There  is  no 
definite  record  of  the  snub-nosed  eel  actually  with- 
in the  southern  rim  of  the  Gulf  so  far  as  we  can 
learn,  and  our  only  first-hand  experience  with  it 
was  on  the  slope  south  of  Nantucket  lightship, 
where  we  captured  21  in  a  Monaco  deep-sea  trap 


'•  The  Japanese  slime  eel,  described  first  as  a  distinct  species  (Uptosomus) 
by  Tanaka  in  190S,  has  been  classed  more  recently  by  him  (Fishes  of  Japan, 
vol.  42, 1928,  p.  810,  pi.  173,  flg,  476)  as  identical  with  the  Atlantic  parasiticu*. 


Figure  71. — Slime  eel  (Simenchelys  parasiticus),  off  Sable  Island  Bank.     From  Goode  and  Bean.     Drawing  by 

H.  L.  Todd. 


158 


FISHERY  BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


in  455  fathoms,  on  the  Grampus  in  July  1908.  It 
must  be  extremely  abundant  along  that  zone,  how- 
ever, for  so  many  to  find  their  way  into  the  trap 
in  as  short  a  set  as  two  hours.  And  it  has  been 
recorded  so  often  in  water  as  shoal  as  200  fathoms 
that  it  may  be  expected  in  the  bottom  of  the 
Eastern  Channel  and  in  the  southeastern  deeps  of 
the  Gulf  of  Maine. 

Long-nosed  eel  Synaphobranchus  pinnatus 
(Gronow)  1854 

Jordan  and  Evermann,  1896-1900,  p.  351. 

Description. — This  deep-sea  species,  a  typical 
eel  in  general  appearance,  is  readily  identifiable 
among  its  tribe  by  the  fact  that  while  its  dorsal 
fin  originates  about  as  far  back  as  in  the  common 
eel  (p.  151),  relative  to  the  length  of  the  fish,  its 
point  of  origin  is  considerably  behind  the  vent 
instead  of  in  front  of  the  latter,  and  that  its  anal 
fin  originates  considerably  in  front  of  the  dorsal 
fin  instead  of  behind  it  as  is  the  case  in  all  other 
Gulf  of  Maine  eels.  Furthermore,  its  mouth  is 
much  wider,  gaping  far  back  of  the  eye,  and  its 
snout  is  pointed.  The  most  interesting  anatomic 
characteristic  of  this  eel  is  that  its  gill  openings, 
opening  longitudinally  on  the  lower  side  of  the 
throat,  join  together  in  front,  apparently  as  a 
single  V-shaped  aperture,  though  actually  they  are 
separate  within. 

Color. — Grayish,  darkest  below,  with  the  ver- 
tical fins  darker  behind  but  pale-edged  in  front, 
and  with  the  inside  of  the  mouth  blue  black. 

Size. — The  largest  of  89  specimens  measured  by 
Goode  and  Bean  was  nearly  22  inches  (545  mm.) 
long,  the  smallest  about  9  inches  (221  mm.)  The 
largest  we  trawled  on  the  Caryn,  in  June  1949, 
was  24  inches  (605  mm.)  long.  Collett "  mentions 
one  26%  inches  (675  mm.)  long  from  the  Azores. 

"  Result,  des  Camp.  Sci.  Prince  de  Monaco,  Pt.  10,  1896,  p.  154. 


Habits. — Nothing  is  known  of  its  habits  except 
that  it  is  a  ground  fish;  that  the  readiness  with 
which  it  bites  a  baited  hook  proves  it  predaceous; 
and  that  specimens  in  spawning  condition  have 
been  taken  in  summer.78  On  June  17,  1949  in 
lat.  42°  38'  N.,  long.  64°  04'  W.,  in  400-460 
fathoms,  we  trawled  many  on  the  Caryn,  both 
males  and  females,  18^  to  24  inches  (470-605 
mm.)  long  that  had  well  developed  gonads,  one 
female  having  already  spawned.  The  ripe  eggs 
are  orange  in  color  and  about  1  mm.  in  diameter. 
In  its  development  it  passes  through  a  lepto- 
cephalus  stage  even  more  slender  than  that  of  the 
American  conger  (p.  156),  and  its  body  segments 
(144-149)  overlap  those  of  the  American  conger 
(140-149)  in  number. 

General  range. — This  deep-water  species  has  a 
wide  distribution.  In  the  western  side  of  the 
North  Atlantic  it  has  been  taken  at  many  local- 
ities along  the  continental  slope  from  the  offing 
of  South  Carolina  to  the  Grand  Banks;  it  is  known 
in  the  east  from  the  Cape  Verdes;  off  Morocco; 
from  the  Canaries;  from  the  Azores;  near  Madeira; 
also  from  the  Faroe  Bank  and  Faroe-Shetland 
Channel.  And  its  leptocephalan  larvae  have  been 
taken  in  such  numbers  from  north  of  Spain  to 
south  of  Iceland  that  it  must  be  one  of  the  most 
plentiful  of  deep-water  fishes  there.79  It  is  also 
recorded  off  Brazil  in  the  South  Atlantic;  likewise 
in  the  Arabian  Sea;  about  the  Philippines;  and  in 
Japanese  waters,  or  is  represented  there  by  a  very 
close  relative.  Most  of  the  captures  have  been 
from  depths  of  300  to  about  2,000  fathoms,  but  it 
has  been  taken  as  shoal  as  129  fathoms. 


"  The  "leptocephalus"  larvae  of  the  long-nosed  eel  axe  described,  with 
photographs  by  Schmidt  (Rapp.  et  Proc.  Verb.  Cons.  Perm.  Intemat. 
Eiplor.  Mer,  vol.  6,  No.  4,  1906,  p.  191,  pi.  9,  flgs.  4-6;  and  Meddel.  Komm- 
Havunderstfgelser,  Ser,  Flskeri,  vol.  3,  No.  6, 1909,  p.  7). 

i»  This  fact  is  commented  on  by  Schmidt  (Rapp.  Cons.  Perm.  Intemat. 
Eiplor.  Mer,  vol.  5,  No.  4,  1906,  p.  191).  For  further  details  as  to  its  distri- 
bution see  Koefoed,  Rept.  Michad  Surj  North  Atlantic  Eiped.,  (1910),  vol. 
4,  Pt.  1, 1927,  pp.  11, 14. 


Figure  72. — Long-nosed  eel  (Synaphobranchus  pinnatus),  La  Have  Bank.     From  Goode  and  Eean.     Drawing  by 

H.  L.  Todd. 


FISHES  OF  THE  GTTLF  OF   MAINE 


159 


Occurrence  in  the  Gulf  of  Maine. — This  eel  has 
not  actually  been  reported  within  the  geographic 
limits  of  the  Gulf.  But  it  is  to  be  expected  in  the 
eastern  channel  and  possibly  above  150  fathoms 
along  the  slopes  of  Georges  Bank,  for  fishermen 
have  caught  them  in  water  as  shallow  as  that  off 
La  Have  Bank,  while  they  have  been  trawled  in 
168  fathoms  and  129  fathoms  off  southern  New 
England  by  the  Fish  Hawk  and  Albatross.  So 
many  of  them  have  been  brought  in  by  fishermen  80 
from  deep  water  off  the  fishing  banks  to  the  east- 
ward of  longitude  65°,  and  so  many  have  been 
trawled  along  the  continental  slope  thence  west- 
ward,81 that  this  eel  must  be  one  of  the  commonest 
of  fishes  below  150  to  200  fathoms,  all  the  way 
from  the  Grand  Banks  to  abreast  of  New  York. 

Snake  eel  Omochelys  cruentifer  (Goode  and  Bean) 
1895 

Jordan  and  Evermann,  1896-1900,  p.  377,  Pisodonophis 
cruentifer. 

Description. — The  most  striking  feature  of  the 
snake  eel  and  one  that  distinguishes  it  from  all 
other  Gulf  of  Maine  eels  is  that  the  tip  of  its  tail 
is  hard  and  pointed.  Other  distinctive  features 
are  that  it  is  only  about  one  thirty-seventh  to 
one  thirty-eighth  as  deep  as  it  is  long;  that  its 
dorsal  fin  originates  only  a  short  distance  behind 
the  tips  of  the  pectorals  when  these  are  laid  back; 
that  its  anal  fin  originates  far  behind  its  dorsal 
fin;  that  its  snout  is  bluntly  pointed;  and  that 
its  mouth  gapes  rearward  considerably  beyond  its 
eyes  (but  not  so  far  back  as  in  the  long-nosed  eel, 
p.  158).  The  dorsal  and  anal  fins  end  a  little  in 
front  of  the  tip  of  the  tail.     The  gill  openings 


*>  Many  such  Instances  are  listed  In  the  Rept.  U.  S.  Comm.  Fish.  (1879) 
1882,  p.  787. 
>>  Goode  and  Bean,  Smithsonian  Contrlb.  Knowl.,  vol.  30, 1895,  pp.  143-144. 


are  short  new-moon-shaped  slits,  close  in  front 
of  the  bases  of  the  pectoral  fins.  Its  "peculiar 
and  savage  physiognomy"  was  stressed  by  its 
describers.82 

Color. — Originally  described  as  uniform  brown- 
ish yellow.  But  those  that  we  have  seen  have 
been  uniform  light  brown  below  as  well  as  above, 
large  ones  darker  than  small  ones.  A  young  one 
about  2%  inches  (6%  cm.)  long  was  pale  with 
dark  speckles. 

Size. — The  largest  yet  seen  was  16%  inches  long. 

Habits. — The  original  account  of  the  snake  eel 
includes  the  information  that  specimens  had  been 
received  that  had  been  taken  from  the  bodies  of 
other  fish,  evidence  that  it  is  a  parasitic-boring 
form.     Nothing  else  is  known  of  its  habits. 

General  range. — Western  side  of  the  Gulf  of 
Maine  to  the  offing  of  Cape  Henry,  Va. 

Occurrence  in  the  Gulf  oj  Maine. — The  snake 
eel  was  originally  described  in  1895  from  6  speci- 
mens trawled  off  Nantucket  by  the  Fish  Hawk, 
and  a  number  have  been  taken  thence  southward 
to  the  latitude  of  Cape  Henry,  Va.,  by  the  Alba- 
tross II,  in  depths  of  24  to  245  fathoms.  The 
only  report  of  it  within  the  Gulf  of  Maine  is  by  its 
describers  of  specimens  taken  by  fishermen  on 
Jeffreys  Bank  many  years  ago. 

Snipe  eel  Nemichthys  scolopaceus  Richardson  1848 

Jordan  and  Evermann,  1896-1900,  p.  369. 

Description. — The  snipe  eel  is  easily  recognizable 
by  its  extremely  slender  body  (the  fish  may  be 
75  times  as  long  as  deep),  with  its  tail  tapering 
to  a  thread,  and  by  its  elongate,  slender,  bill-like 
jaws,  one  as  long  as  the  other,  the  upper  one 
curving    upward,    but    the    lower    more    nearly 


"  Goode  and  Bean,  Smithsonian  Contrlb.  Knowl.,  vol.  30, 1895,  p.  147. 


Figure  73. — Snake  eel  (Omochelys  cruentifer),  continental  slope  south  of  Nantucket  Shoals.     From  Goode  and  Bean. 

Drawing  by  H.  L.  Todd. 


160 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE    SERVICE 


2^22222222222^2; 


>a^""""v»"^vi^w>>Y.^^^^^^^^SSi^^ 


Figure  74. — Snipe  eel  (Nemichthys  scolopaceus) .     Drawing  by  E.  N.  Fischer,  from  Bigelow  and  Welsh. 


straight.  The  head  is  much  deeper  than  the 
neck,  with  large  eyes.  The  dorsal  fin  originates 
in  front  of  the  pectorals,  the  anal  about  abreast 
of  the  tip  of  the  pectorals,  and  both  dorsal  and 
anal  run  back  to  the  tip  of  the  tail. 

There  has  been  some  confusion  in  the  published 
accounts  and  illustrations  as  to  the  dorsal  and 
anal  fins,  for  while  Vaillant83  shows  both  as  about 
as  high  throughout  their  length  as  the  fish  is  deep, 
Goode  and  Bean84  picture  the  dorsal  as  much 
higher  than  the  anal  (the  artist  evidently  having 
transposed  the  two  fins) ,  whereas  Brauer 86  repre- 
sents the  anal  as  approximately  twice  as  high  as 
the  dorsal  and  the  latter  as  soft  rayed  in  its  an- 
terior and  posterior  portions  but  composed  of 
short  thorn-like  spines  along  its  central  third.  The 
fins  of  two  specimens  taken  off  New  England,  now 
in  the  collections  of  the  Museum  of  Comparative 
Zoology  are  as  follows: 

Dorsal,  soft-rayed  and  nearly  as  high  as  the 
body  is  deep  for  its  first  half;  back  of  that  it  con- 
sists of  a  series  of  very  short,  stiff  rays  that  extend 
to  the  tip  of  the  tail. 

Anal,  soft-rayed  throughout  its  length  and  about 
as  high  as  the  body  is  deep,  tapering  to  almost 
nothing  on  the  tail. 

The  confusion  has  been  due  in  part  to  the  rather 
fragmentary  state  in  which  these  deep-water  fish 
usually  arrive  on  board,  but  at  the  same  time  it  is 
probable  that  two  distinct  species  have  been  con- 
fused under  the  name  scolopaceus,  as  Brauer 
suspected. 


Color. — Described  as  pale  to  dark  brown  above 
with  the  belly  and  anal  fin  blackish  after  preserva- 
tion. Judging  from  experience  with  other  deep- 
sea  fishes  and  from  Brauer's  plate  (which,  however, 
may  be  another  species),  we  suspect  that  it  is 
chocolate  brown  above  in  life  and  velvety  black 
below. 

Size. — Maximum  length  about  3  feet. 

Habits. — Although  commonly  spoken  of  as  a 
"deep-sea"  fish,  this  species  is  undoubtedly  an 
inhabitant  of  the  mid  depths,  not  of  the  bottom, 
and  judging  from  the  occurrence  of  other  black 
fishes  it  probably  finds  its  upper  limit  at  100  to 
200  fathoms.  Nothing  further  is  known  of  its 
habits,  but  Mowbray's 86  capture  near  Bermuda  of 
a  snipe  eel  clinging  by  its  jaws  to  the  tail  of  a  large 
red  snapper  has  suggested  that  such  may  be  a 
regular  habit  of  this  curious  species. 

General  range. — The  snipe  eel  has  been  taken 
in  deep  water  at  many  stations  off  the  east  coast 
of  North  America  between  latitudes  31°  and  42°N., 
longitudes  65°  and  75°W. ;  also  in  the  South  Atlan- 
tic; near  the  Azores;  near  Madeira;  off  the  Cape 
Verde  Islands ;  off  West  Africa ;  and  in  the  Pacific 
of  New  Guinea. 

Occurrence  in  the  Gulf  of  Maine. — One  specimen 
taken  from  the  stomach  of  a  codfish  caught  on 
Georges  Bank  in  45  fathoms  is  the  only  Gulf  of 
Maine  record,  but  several  have  been  taken  in 
depths  of  from  300  to  2,000  fathoms  on  the  sea- 
ward slope  of  the  bank. 


THE  LANCET  FISHES.     FAMILY  ALEPISAURIDAE 


The  lancet  fishes  have  one  long  and  very  high 
dorsal  fin,  soft-rayed  from  end  to  end;  a  small 

-  Poissons.  Exped.  Scl.  Travailleur  et  Talisman,  1888,  pi.  7,  figs.  2  and  2a. 
M  Smithsonian  Contrlb.  Knowl.,  vol.  31,  1895,  pi.  46,  fig.  170. 
'« Tiefsee-Flsche,  Wlss.  Ergeb.  Deutsch.  Tiefsee-Exped.  (189S-1899),  vol 
16,  Pt.  1, 1900,  p.  126,  pi.  9,  fig.  1. 


adipose  fin  behind  the  dorsal  fin,  like  that  of  a 
salmon  or  smelt;  a  deeply  forked  caudal  fin;  a 
short  anal,  most  of  which  is  behind  the  rear  end 
of  the  dorsal;  large  pointed  pectorals  and  ven- 


"Copela  ,No.  108. 1922,  p.  49. 


FISHES   OF  THE  GULF  OF   MAINE 


161 


trals;  and  a  very  wide  mouth  with  large  teeth. 
Several  species  are  known,  all  belonging  to  deep 
water;  only  one  has  been  taken  within  the  province 
covered  by  this  report.  Their  closest  affinities 
seem  to  lie  with  the  lanternfishes  (p.  141). 

Lancetfish  Alepisaurus  ferox  Lowe  1833 
Handsawfish 

Jordan  and  Evermann,  1896-1900,  p.  595. 

Description. — The  combination  of  a  long  and 
very  high  dorsal  fin,  soft-rayed  from  end  to  end, 
with  the  presence  of  an  adipose  fin  behind  it, 
distinguishes  the  lancetfish  from  all  other  Gulf 
of  Maine  fishes.  The  body  is  slender,  somewhat 
flattened  sidewise,  deepest  at  the  gill  covers,  and 
tapers  back  to  a  slender  caudal  peduncle.  The 
snout  is  long  and  pointed,  the  mouth  wide, 
gaping  back  of  the  eye,  and  each  jaw  has  two 
or  three  large  fangs,  besides  smaller  teeth.  The 
dorsal  fin  (41  to  44  rays)  originates  on  the  nape 
and  occupies  the  greater  length  of  the  back,  is 
rounded  in  outline,  about  twice  as  high  as  the 
fish  is  deep,  and  can  be  depressed  in  a  groove 
along  the  back.  The  adipose  fin  recalls  that  of 
the  smelt  in  form  and  location.  The  caudal  is 
very  deeply  forked;  its  upper  lobe  is  prolonged  as 
a  long  filament,  and  although  most  of  the  speci- 
mens so  far  seen  have  lost  this  we  have  an  ex- 
cellent photograph  showing  it.  The  anal  fin 
originates  under  the  last  dorsal  ray,  and  is  deeply 
concave  in  outline.  The  ventrals  are  about 
halfway  between   the   anal   and   the   tip   of  the 


snout,  while  the  pectorals  are  considerably  longer 
than  the  body  is  deep  and  are  situated  very  low 
down  on  the  sides.  There  are  no  scales  and  the 
fins  are  exceedingly  fragile. 

Color. — Sides  described  as  metallic  silvery.  We 
have  not  seen  a  newly  taken  specimen. 

Size. — The  collection  of  the  Boston  Society  of 
Natural  History  contains  the  cast  of  a  specimen 
about  6  feet  long  that  was  taken  off  Nova  Scotia 
in  August  1910,  and  this  is  probably  about  the 
maximum  size. 

Habits. — This  is  an  oceanic  species,  of  the 
mid-depths,  appearing  only  as  a  stray  shoaler 
than  200  fathoms.  Nothing  is  known  of  its 
habits.  A  Block  Island  specimen  had  eaten  a 
small  spiny  dogfish. 

General  range. — Widely  distributed  in  the  deep 
waters  of  the  Atlantic,  also  reported  from  the 
northeastern  Pacific.87 

Occurence  in  the  Gulf  oj  Maine. — A  specimen 
brought  in  by  a  fisherman  from  Georges  Bank  88 
about  1878  or  1879  is  its  only  claim  to  mention 
here.  Goode  and  Bean  and  Vladykov  and 
McKenzie 89  have  reported  other  captures  of  lan- 
cetfishes  from  La  Have  Bank,  from  southeast  of 
Emerald  Bank  and  Banquereau.  Another  speci- 
men 5%  feet  long  was  caught  alive  in  the  surf  on 
Block  Island,  R.  I.,  March  12,  1928,  and  reported 
by  Mrs.  Elizabeth  Dickins  who  sent  us  a  photo- 
graph of  it. 


"  Crawford  (Copela,  No.  164,  1927,  p.  66)  reports  several  A.  ferox  from 
the  halibut  banks  off  the  northwestern  coast  of  British  Columbia. 
M  No  definite  information  is  available  as  to  this  specimen. 
'•  Proc.  Nova  Scotia  Inst.  Sci.,  vol.  19, 1935,  p.  63. 


Figube  75. — Lancetfish  (Alepisaurus  ferox) .     New  York  market  specimen.     From  Jordan  and  Evermann.     Drawing  by 

H.  L.  Todd. 
210941—53 12 


162 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


THE  MUMMICHOGS  OR  KILLIFISHES.  FAMILY  POECILIIDAE 


The  mummichogs  are  small  fishes  made  rec- 
ognizable by  having  only  one  short  soft-rayed 
dorsal  fin  situated  far  back,  and  ventrals  situ- 
ated on  the  abdomen,  combined  with  a  small 
mouth  at  the  tip  of  the  snout,  a  very  thick  caudal 
peduncle,  and  a  rounded  tail  fin.  The  family  is 
represented  in  the  Gulf  of  Maine  by  three  species, 
two  of  Funduius  and  one  of  Cyprinodon,  the  former 
slender  and  the  latter  deep  in  outline,  a  dif- 
ference in  body  form  sufficient  to  distinguish  the 
one  genus  from  the  other  at  a  glance.  The 
teeth  are  likewise  different  in  the  two  genera, 
those  of  Fundvlus  being  sharp-pointed,  whereas 
they  are  wedge-shaped  in  Cyprinodon  and  in- 
cisorlike. The  two  local  species  of  Funduius 
are  separable  by  their  markings,  majalis  of  all 
ages  being  barred  or  streaked  with  black  while 
the  adult  heteroclitus  is  not. 

Common  mummichog  Funduius  heteroclitus 
(Linnaeus)  1766 

Killifish;  Salt-water  minnow;  Chub;  Mummy 

Jordan  and  Evermann,  1896-1900,  p.  640. 

Description. — This  is  a  stout-bodied  little  fish, 
about  one-fourth  as  deep  as  long,  its  body  thickest 
just  back  of  the  pectoral  fins,  whence  it  tapers  to 
the  tail.  Both  its  back  and  its  belly  are  rounded, 
but  the  top  of  the  head  is  flat  between  the  eyes. 
The  snout,  as  seen  from  above,  is  blunt.  The 
mouth  is  at  the  tip  of  the  snout  and  is  so  small  that 
it  does  not  gape  back  to  the  eye.  Perhaps  the  most 
striking  feature  of  Funduius  is  its  very  deep  caudal 


peduncle  and  rounded  caudal  fin.  The  fins  are  of 
moderate  size,  the  dorsal  situated  behind  the 
middle  of  the  body  above  the  anal,  the  pectorals 
broad  and  rounded.  Both  head  and  body  are 
covered  with  large  rounded  scales.  On  males  in 
breeding  condition  the  scales  on  the  sides  of  the 
head  and  those  on  the  flanks  below  and  behind  the 
dorsal  fin  develop  fingerlike  processes  on  their  free 
edges,  called  "contact  organs." 

The  mummichog  shows  a  striking  sexual  dimor- 
phism in  the  dorsal  and  anal  fins,  which  are  not 
only  larger  in  the  male  than  in  the  female,  and  the 
anals  of  a  different  shape,90  but  are  more  muscular 
and  are  used  as  claspers  in  the  act  of  spawning. 

Color. — Males  and  females  differ  in  color  as  well 
as  in  the  sizes  of  the  fins.  Out  of  breeding  season 
the  males  are  dark  greenish  or  steel  blue  above, 
with  white  and  yellow  spots,  and  marked  on  the 
sides  with  narrow  irregular  silvery  bars  or  mot- 
tlings  made  up  a  series  of  dots.  The  belly  is 
white,  pale  yellow,  or  orange;  the  dorsal,  anal,  and 
caudal  fins  are  dark  green  or  dusky  with  pale 
mottlings;  the  front  edges  of  the  anal  and  of  the 
ventrals  are  yellow.  Sometimes  there  is  a  dark- 
edged,  pale-centered  eyespot  on  the  rear  part  of 
the  dorsal  fin.  At  spawning  time  the  pigmenta- 
tion of  the  male  is  generally  intensified,  the  back 
and  upper  sides  darkening  almost  to  black,  while 
the  yellow  of  the  belly  becomes  more  brilliant  and 
the  body  generally  takes  on  steel-blue  reflections. 
The  females  (much  paler  than  the  males)  are  uni- 


"  A  detailed  account  of  the  sexual  differences  Is  given  by  Newman  (Biol. 
Bull.,  vol.  12,  No.  5, 1907,  pp.  314-348). 


Figure  76. — Common  mummichog   (Funduius  heterocliius),   Maryland. 

A.  H.  Baldwin. 


From  Jordan  and   Evermann.     Drawing  by 


FISHES   OF  THE  GULF  OF   MAINE 


163 


form  olive  to  bottle  green,  darker  above,  lighter 
below,  without  definite  markings  though  their 
sides  often  show  faint  and  indefinite  crossbars  of  a 
deeper  tone  of  the  same  hue.  Their  fins  are  much 
paler  than  those  of  males.  Very  young  fry  of 
both  sexes  show  dark  transverse  bars  on  the  sides, 
but  these  bars  are  lost  with  growth. 

Killifishes  vary  in  shade  from  very  pale  to  dark, 
according  to  the  color  of  their  surroundings.  And 
recent  experiments 9l  have  proved  that  their 
ability  to  change  from  light  to  dark  depends  on  the 
sense  of  sight. 

Size. — The  maximum  length  is  5  to  6  inches, 
but  adult  mummichogs  are  seldom  more  than  3K 
to  4  inches  long  in  the  Gulf  of  Maine.  Several 
varieties  of  this  species  have  been  described,  but 
they  are  so  closely  allied  that  it  is  not  necessary  to 
discuss  them  here. 

Habits. — The  home  of  the  mummichog  is  along 
sheltered  shores  where  the  tide  flows  over  beds  of 
eelgrass  or  salt  hay  (Spartina),  among  which  shoals 
of  "mummies"  may  often  be  seen  moving  in  with 
the  flood  tide.  They  abound  in  the  tidal  creeks 
that  cut  the  salt  marshes,  on  the  shores  of  our 
harbors,  and  in  the  brackish  water  at  the  mouths 
of  our  streams  and  estuaries,  particularly  in  little 
muddy  pools,  creeks,  and  ditches.  So  closely, 
indeed,  do  they  hug  the  shore  that  a  line  drawn 
100  yards  out  from  land  would  probably  inclose 
practically  all  the  mummichogs  in  the  Gulf  of 
Maine.  Where  the  shore  is  bold  and  rocky,  as  it  is 
about  the  Bay  of  Fundy,  the  mummichog  is 
practically  restricted  to  brackish  water,  and  it 
often  goes  up  into  fresh  water.  At  the  other 
extreme,  it  is  not  likely  that  they  ever  descend  to 
a  depth  of  more  than  a  couple  of  fathoms  in  their 
journeys  in  and  out  of  the  creeks  or  along  the 
shore. 

Mummichogs  are  so  resistant  to  a  lack  of  oxy- 
gen, to  the  presence  of  carbon  dioxide,  and  to  un- 
favorable surroundings  generally,  that  they  can 
survive  in  very  foul  water.  At  ebb  tide  "mum- 
mies" are  often  trapped  in  little  pools  where  they 
remain  until  the  next  tide  if  the  water  holds,  often 
huddled  together  in  swarms.  Should  the  pool  go 
dry,  they  work  their  way  for  the  time  being  into 
the  mud,  where  we  have  often  found  them.02  And 
it  is  probable  that  they  can  flop  overland  for  a  few 


yards  to  some  other  drain  as  the  striped  mummi- 
chog does. 

They  winter  in  a  more  or  less  sluggish  state  on 
the  bottoms  of  the  deeper  holes  or  creeks,  where 
they  have  been  found  buried  6  or  8  inches  deep  in 
the  mud,93  and  there  is  no  evidence  that  they  move 
out  to  sea  during  the  cold  season;  in  short,  this  is 
one  of  the  most  stationary  of  fishes. 

Food. — "Mummies"  are  omnivorous,  feeding  on 
all  sorts  of  edible  things,  vegetable  as  well  as 
animal.  They  have  been  found  full  of  diatoms, 
eelgrass,  and  other  vegetable  matter;  foraminif- 
era;  shrimps  and  other  small  Crustacea,  also 
small  mollusks;  and  they  sometimes  have  small 
fish  in  their  stomachs.94 

They  soon  congregate  about  any  dead  fish  or 
other  bit  of  carrion,  to  prey  either  upon  it  or  upon 
the  amphipod  scavengers  that  gather  on  such 
dainties  and  they  eat  their  own  or  each  other's 
eggs  at  spawning  time. 

Spawning  probably  takes  place  at  the  same 
season  in  the  Gulf  of  Maine  as  on  the  southern 
coast  of  New  England;  i.  e.,  in  June,  July,  and 
early  August.  As  sexual  activity  approaches,  the 
males  (now  brilliantly  tinted)  court  and  pursue 
the  females,  rivalry  among  them  being  very  keen; 
those  that  are  the  most  highly  colored  or  most 
excited  usually  driving  off  the  others.  Sometimes 
they  fight  fiercely.  They  spawn  in  a  few  inches  of 
water,  seeking  shady  spots.  At  the  moment  of 
spawning  the  male  clasps  the  female  with  his  anal 
and  dorsal  fins  just  back  of  her  anal  and  dorsal, 
usually  forcing  her  against  some  stone  or  against 
the  bottom,  the  bodies  of  both  are  bent  into  an  S 
and  their  tails  vibrate  rapidly  while  the  eggs  and 
the  milt  are  extruded.95  Occasionally,  pairs  clasp 
and  spawn  free  in  the  water  without  coming  in 
contact  with  any  object,  and  sometimes  a  female 
is  seen  to  pursue  and  court  a  male. 

The  eggs,  which  are  about  2  mm.  in  diameter, 
colorless  or  pale  yellowish  and  surrounded  by  a 
firm  capsule,  sink  and  become  so  sticky  on  contact 
with  the  water  that  they  mass  together  in  clumps, 
or  stick  fast  to  sand  grains  or  to  anything  else 
they  chance  to  rest  upon.  Incubation  occupies 
from  9  to  18  days,  the  exact  duration  probably 


•'  Parker  and  Lanchner,  Amer.  Jour,  of  Physiol.,  vol.  61, 1922,  p.  648. 
••  This  habit  Is  described  by  Mast  (Jour,  of  Animal  Behavior,  vol.  5,  No.  5, 
1915,  p.  351). 


•I  Chichester,  Amer.  Naturalist,  vol.  64, 1920,  p.  651. 

•<  Lists  of  stomach  contents  are  given  by  Field  (Rept.  U.  S.  Comm.  Fish., 
H906)  1907,  p.  29). 

«  Newman  (Biol.  Bull.,  vol.  12,  No.  5,  April  1907,  p.  315)  gives  an  Interest- 
ing account  of  the  courtship  and  spawning,  from  which  the  preceding  la 
condensed. 


164 


FISHERY  BULLETIN    OF   THE    FISH   AND   WILDLIFE    SERVICE 


depending  on  temperature.  The  larva  is  about  7  to 
7.7  mm.  long  at  hatching,  its  yolk  absorbed  al- 
ready, and  its  pectoral  and  caudal  fins  fully  formed. 
By  the  time  the  little  fish  has  grown  to  11  mm.  the 
dorsal  and  anal  fin  rays  are  present  in  full  number, 
and  the  first  trace  of  the  ventrals  is  to  be  seen.  At 
16  mm.  the  ventrals  are  apparent,  and  fry  of  20 
mm.  resemble  their  parents. 

General  range. — Coast  of  North  America,  from 
the  Gulf  of  St.  Lawrence  to  Texas.  Port  au  Port 
Bay,  on  the  west  coast  of  Newfoundland,95  is  the 
most  northerly  record  that  we  have  found  for  it. 

Occurrence  %n  the  Gulf  of  Maine. — The ' '  mummie' ' 
is  one  of  the  few  fish  which  can  fairly  be  charac- 
terized as  "universal"  in  suitable  locations  around 
the  entire  coastline  of  the  Gulf.  We  dare  say 
that  there  is  not  a  single  bit  of  salt  marsh,  muddy 
creek,  harbor,  sheltered  shore  line,  or  brackish 
estuary,  where  they  are  not  to  be  found,  from  the 
elbow  of  Cape  Cod  around  to  Cape  Sable. 


••  Johansen  (Canadian  Naturalist,  vol.  40,  February  1926,  p.  34). 


Importance. — The  mummichog  is  of  some  com- 
mercial value  as  bait,  but  only  locally.  It  is 
also  a  favorite  for  biological  experiment. 

Striped   mummichog  Fundulus   majalis    (Wal- 

baum)  1792 

Mummichog;  Mummy;  Killifish 

Jordan  and  Evermann,  1896-1900,  p.  637. 

Description. — This  fish  resembles  the  common 
mummichog  closely  in  general  form,  in  the  shape 
of  its  dorsal  and  aDal  fins,  in  its  sexual  dimorphism, 
and  in  the  development  of  "contact  organs"  on  the 
scales  of  breeding  males.  But  it  is  more  slender, 
its  snout  more  pointed,  its  body  more  definitely' 
fusiform,  tapering  toward  both  head  and  tail, 
and  its  caudal  peduncle  not  so  stout.  But  the 
color  pattern  is  the  most  striking  difference 
between  the  two,  both  sexes  of  Fundulus  majalis 
being  definitely  barred  with  black  at  maturity 
as  well  as  when  young.  In  the  male  the  barring 
is  transverse  throughout  life,  the  stripes  increasing 


Figure  77. — Striped  mummichog  (Fundulus  majalis).     Upper,  male,  Woods  Hole;  lower,  female,   Maryland. 

Jordan  and  Evermann. 


From 


FISHES   OF  THE  GULF  OF   MAINE 


165 


in  number  from  7  to  12  in  the  young  to  14  to  20  in 
adult  fish.  When  the  females  reach  a  length  of 
about  two  inches,  however,  the  original  7  to  12 
transverse  bars  are  transformed  with  growth  into 
two  or  three  longitudinal  stripes  on  each  side,  the 
upper  stripe  running  uninterrupted  from  gill 
opening  to  tail,  the  lower  stripes  in  two  segments, 
the  one  from  close  behind  the  pectoral  to  above 
the  ventral,  the  other  thence  backward  to  close 
behind  the  rear  edge  of  the  anal  fin.  One  or  two 
transverse  bars  persist  however  on  the  caudal 
peduncle,  even  on  the  oldest  females. 

Color. — This  is  a  decidedly  paler  fish  than  the 
other  "mummy."  Apart  from  the  black  bars 
the  male  is  dark  olive  green  above  with  silvery 
sides,  a  greenish-yellow  belly,  and  a  black  spot 
on  the  rear  part  of  the  dorsal  fin;  his  pectorals  and 
caudal  are  pale  yellowish.  The  male  becomes 
more  brilliant  at  breeding  time,  the  back  turning 
almost  black,  the  lower  sides  and  belly  changing  to 
orange  or  golden,  and  the  fins  to  bright  yellow. 
The  female  is  olive  green  above  and  white  below, 
striped  as  described  above. 

Size. — This  is  a  larger  fish  than  the  common 
mummichog,  occasionally  growing  to  a  length  of 
7  inches  and  often  to  6  inches. 

Habits. — The  striped  mummie  parallels  the 
common  mummie  in  being  restricted  to  the 
immediate  neighborhood  of  the  land,  and  in  its 
way  of  life,  except  that  it  keeps  more  strictly  to 
salt  water,  and  is  found  more  often  along  open 
beaches.  Its  most  interesting  habit  is  its  ability 
to  flop  back  into  the  water  if  it  becomes  stranded 
with  the  receding  tide,  jumping  unerringly  toward 
the  water  in  almost  every  instance,  and  progressing 
from  several  inches  to  several  feet  at  each  jump.97 
And  so  noted  are  they  for  this  peculiar  ability 
that  a  special  article  has  been  devoted  to  it.88 
Their  diet  consists  of  small  animals  including 
mollusks,  crustaceans,  fish,  insects,  and  insect 
larvae.  Westward  and  southward  from  Cape 
Cod  they  spawn  from  late  spring  to  late  summer. 

General  range. — Coast  of  the  United  States, 
from  the  vicinity   of  Boston,  Mass.,  to  Florida. 

Occurrence  in  the  Gulf  of  Maine. — The  striped 
mummie  is  very  abundant  along  the  southern 
shores  of  New  England,  westward  from  Cape 
Cod.     But  the  only  published  records  for  it  in 


the  gulf  are  for  the  vicinity  of  Boston  and  Salem, 
many  years  ago,  and  we  had  not  seen  it  north  of 
Cape  Cod  before  1937.  In  that  autumn,  how- 
ever, B.  Preston  Clark  brought  in  four  specimens 
that  he  had  taken  at  Cohasset,  on  the  southern 
shore  of  Massachusetts  Bay;  it  was  reported  to 
us  as  in  numbers  there  in  1939,"  and  we  have 
seen  small  schools  of  them  in  recent  summers 
in  the  salt  marshes  at  the  entrance  to  Cohasset 
Harbor,  as  well  as  nearby.  If  this  little  fish 
actually  has  extended  its  regular  range  north- 
ward and  if  its  dispersal-route  has  been  via  the 
Cape  Cod  Canal,  as  has  been  suggested,1  it  is  to  be 
expected  anywhere  in  the  marshes  around  Cape 
Cod  Bay  and  along  the  southern  shore  of  Massa- 
chusetts Bay,  and  we  suspect  that  a  resident 
population  is  to  be  found  in  the  Nauset  Marshes 
and  in  Pleasant  Bay,  on  the  outer  shore  of  Cape 
Cod. 

Sheepshead      minnow     Cyprinodon    variegatus 
Lacep&de  1803 

Jordan  and  Evermann,  1896-1900,  p.  671. 

Description. —  The  sheepshead  minnow  is  so 
deep  bodied  (its  body  is  nearly  half  as  deep  as 
long,  not  counting  the  tail  fin)  that  it  is  not  apt 
to  be  mistaken  for  either  of  the  mummichogs. 
And  it  is  separated  further  from  the  mummi- 
chogs by  its  teeth,  which  are  large,  wedge-shaped 
and  with  tricuspid  cutting  edges,  instead  of  small 
and  pointed.  It  is  a  flat-sided  little  fish,  with 
high  arched  back,  small  flat-topped  head,  small 
terminal  mouth  hardly  gaping  back  to  the  forward 
edge  of  the  eye,  and  it  has  the  thick  caudal 
peduncle  characteristic  of  its  family.  Its  tail  is 
square  (rounded  in  the  mummichogs),  and  the 
fact  that  almost  the  whole  of  its  dorsal  fin  is  in 
front  of  the  anal  instead  of  over  it  affords  an- 
other point  of  difference.  The  pectorals  are 
large,  reaching  back  past  the  base  of  the  ventrals, 
which  seem  very  small  by  contrast.  Both  its 
body  and  its  head  are  covered  with  large  rounded 
scales,  largest  on  top  of  the  head  and  on  the 
cheeks,  with  one  much  larger  than  the  others  just 
above  the  pectoral  fin.  Young  fish  are  propor- 
tionally more  slender  than  old  ones.  The  dorsal, 
ventral,  and  anal  fins  are  higher  in  the  males  of 
this  species  than  in  the  females,  much  as  they 


•'  Hildebrand  and  Schroeder,  Bull.  U.  8.  Bur.  Fish.  vol.  48,  Pt.  1    1928 
p.  141. 
•'  Mast,  Jour  .o  (Anlma  Behavior,  vol.  5,  No.  5, 1916,  pp.  341-340. 


«  By  John  W.  Lowes. 

i  Schroeder,  Copela,  1937,  No.  4,  p.  238. 


166 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure   78. — Sheepshead   minnow    (Cyprinodon   variegatus),    Maryland.     From   Jordan   and   Evermann.     Drawing   by 

A.  H.  Baldwin. 


are  in  the  mummichogs.  The  male  is  deeper 
bodied  and  the  average  size  is  larger  than  that 
of  the  female. 

Color. — Out  of  breeding  season  both  males  and 
females  are  olive  above  (males  rather  darker  and 
greener  than  females)  with  pale  yellow  or  yellow- 
ish-white belly,  dusky  dorsal  fin,  and  pale  orange 
pectoral,  ventral,  and  anal  fins.  The  young  of 
both  sexes  are  irregularly  barred  with  black 
transverse  stripes,  which  persist  through  life  in 
the  female  but  become  obscured  in  adult  males. 
Females,  furthermore,  have  a  black  spot  on  the 
rear  corner  of  the  dorsal  fin,  which  is  lacking  in 
males,  while  the  caudal  fin  of  the  male  is  marked 
by  two  black  cross  stripes,  one  at  the  base  and 
the  other  at  the  margin.  In  breeding  season 
the  male  assumes  a  brilliant  coat,  his  upper 
parts  turning  to  steel  blue  in  front  of  the  dorsal 
fin  with  a  greenish  luster  behind  it,  while  his 
belly  brightens  to  a  deep  salmon,  his  ventrals 
and  anal  change  to  dusky  margined  with  orange, 
and  the  front  edge  of  his  dorsal  turns  orange. 

Size.— The  largest  specimens  are  about  3 
inches  long. 

Habits. — The  sheepshead  minnow  (like  the  com- 
mon mummichog)  is  confined  to  the  shallow  waters 
of  inlets,  harbors,  and  the  heads  of  bays,  and  salt 
marshes,   often   in   brackish   water.     Its   diet   is 


partly  vegetable,  partly  animal.  It  is  very  pug- 
nacious, often  killing  fishes  larger  than  itself, 
making  repeated  attacks  with  its  sharp  teeth  and 
finally  devouring  its  victim.  Its  breeding  habits 
recall  those  of  the  mummichog  (p.  163),  the  males 
fighting  fiercely  among  themselves  and  clasping 
the  females  just  forward  of  the  tail  with  dorsal 
and  anal  fin,  while  the  eggs  and  milt  are  extruded. 
Spawning  takes  place  in  shallow  water  from  April 
to  September,  the  eggs  maturing  a  few  at  a  time, 
so  that  any  given  female  spawns  at  intervals 
throughout  the  season.  The  eggs  sink  and  stick 
together  in  clumps  by  numerous  threads.  They 
are  1.2  to  1.4  mm.  in  diameter,  with  one  large  oil 
globule  and  many  minute  ones.  Incubation  oc- 
cupies 5  or  6  days,  and  even  at  hatching  the  larvae 
(4  mm.  long)  show  alternate  light  and  dark  cross- 
bands.  At  a  length  of  9  mm.  all  the  fins  are 
formed,  and  at  12  mm.  the  fry  show  most  of  the 
characters  of  its  parents.2 

General  range. — Atlantic  coast  of  the  United 
States,  Cape  Cod  to  Mexico,  in  brackish  as  well 
as  in  salt  water. 

Occurrence  in  the  Gulf  of  Maine. — This  fish,  like 
many  others,  finds  its  northern  limit  at  Cape  Cod 


1  An  account  of  courtship  and  spawning  is  given  by  Newman  (Biol.  Bull., 
vol.  12,  No.  8, 1907,  p.  336)  and  of  development  by  Kuntz  (Bull.,  U.  S.  Bur. 
of  Fish.,  vol.  34,  (1914)  1916,  p.  409). 


FISHES   OF  THE  GULF  OF   MAINE 


167 


and  would  not  deserve  mention  here  at  all  were  it 
not  recorded  from  the  Cape  by  Storer.  West  and 
south  of  Cape  Cod,  however,  as  at  Woods  Hole, 


it  is  common  enough  in  suitable  situations.  We 
have  seined  many  of  them  with  Fundvlus  at  the 
head  of  Buzzards  Bay. 


THE  BILLFISHES  OR  SILVER  GARS.     FAMILY  BELONIDAE 


The  most  noticeable  feature  of  the  billfishes  is 
that  both  of  their  jaws  are  prolonged  to  form  a  long 
slim  beak  well  armed  with  teeth.  Their  bodies 
are  very  slender,  and  their  anal,  dorsal,  and  ventral 
fins  set  far  back.  They  have  no  finlets  between 
the  dorsal  and  anal  fins  and  the  caudal,  the  absence 
of  these  being  the  readiest  field  mark  to  separate 
the  billfishes  from  the  needlefishes  (Scomber esox,  p. 
170).  They  are  swift-swimming,  predaceous  fishes, 
represented  by  many  species,  most  of  them 
American.  Only  two  have  ever  been  recorded  in 
the  Gulf  of  Maine.3 

Key  to  Gulf  of  Maine  Billfishes 

Body  as  thick  as  it  is  deep;  dorsal,  anal,  and  caudal  fins 
only  moderately  concave Silver  Gar,  p.  167 

Body  less  than  }i  as  thick  as  it  is  deep;  dorsal,  anal,  and 
caudal  fins  deeply  concave Garfish,  p.  168 

Silver  gar  Tylosurus  marinus  (Walbaum)  1792  4 

Billfish;  Salt-water  gar;  Sea  pike;  and 
various  other  local  names 

Jordan  and  Evermann,  1896-1900,  p.  714. 

Description. — Its  long  bill  and  slender  body  give 
the  gar  so  peculiar  an  aspect  that  it  is  not  likely  to 
be  confused  with  any  other  Gulf  of  Maine  fish 


'The  closely  allied  houndflsh  (Tylosurus  acus  Lacepe.de  1803)  has  been 
taken  at  Nantucket,  but  has  not  been  found  within  the  Oulf  of  Maine. 
Since  It  may  appear  there  as  a  stray  from  the  south  ,  we  may  point  out  that  it 
Is  easily  distinguished  from  the  sliver  gar  by  Its  de  eply  forked  tall  and  by  the 
fact  that  Its  dorsal  and  anal  fins  are  much  longer,  the  former  with  23  rays,  the 
latter  with  21.  The  following  characters  In  combination  will  serve  to  Identify 
It  among  the  several  tropical  gars;  mouth  nearly  closable  and  upper  jaw  not 
arched;  dorsal  and  anal  fins  long;  beak  at  least  twice  as  long  as  rest  of  head; 
greatest  depth  of  body  not  more  than  two-thirds  as  great  as  length  of  pec- 
toral fin;  no  lateral  stripe. 

•  Jo  rdan,  Evermann,  and  Clark  (Kept.  U.  8.  Comm.  Fish.,  (1928)  Pt.  2, 
1930,  p.  196)  place  this  species  In  the  genus  Strongylum  Van  Hasselt  1824. 


other  than  the  half  beak  (p.  169),  the  needle  fish 
(p.  170),  or  its  own  close  relative  Ablennes  (p.  168). 
And  it  is  easily  distinguishable  from  the  first  of 
these  by  the  fact  that  both  of  its  jaws  are  prolonged 
instead  of  only  the  lower;  from  the  second  by  lack- 
ing detached  finlets  on  its  back  between  its  dorsal 
and  caudal  fins.  The  most  conspicuous  differ- 
ences between  the  silver  gar  and  Ablennes  (p.  168) 
is  that  the  body  of  the  former  is  thicker  than  it  is 
deep,  and  that  its  fins  are  only  moderately  con- 
cave, whereas  the  latter  is  so  strongly  flattened 
sidewise  that  it  is  less  than  one-half  as  thick  as  it 
is  deep  with  deeply  concave  fins. 

The  head  of  the  adult  silver  gar  occupies  nearly 
one-third  of  the  total  length  of  the  fish;  the  upper 
jaw,  from  the  eye  forward,  is  twice  as  long  as  the 
rest  of  the  head;  both  jaws  are  armed  with  sharp 
teeth;  and  the  eyes  are  large.  The  long,  slender 
body  is  only  about  one-twentieth  as  deep  as  long, 
rounded  (not  laterally  flattened)  in  cross  section, 
and  thicker  than  deep.  Both  the  body  and  the 
sides  of  the  head  are  scaly.  The  dorsal  fin,  with 
13  to  17  rays,  and  the  anal  fin,  with  17  to  21  rays, 
are  alike  in  outline,  the  anterior  rays  of  both  being 
much  longer  than  those  toward  the  rear,  and  the 
rear  two-thirds  of  each  can  be  depressed  along  the 
back  and  nearly  concealed  in  a  groove,  while  the 
forward  one-third  continues  erect.  Both  fins,  too, 
are  situated  far  back,  with  the  dorsal  arising  a 
little  behind  the  forward  end  of  the  anal. 

The  ventral  fins  stand  about  halfway  between 
a  point  below  the  eye  and  the  base  of  the  caudal. 
The  margin  of  the  caudal  fin  is  only  moderately 
concave,  this  fact  being  the  readiest  field  mark  to 
separate  this  particular  gar  from  the  only  other 
species  of  its  genus  (Tylosurus  acus)  taken  yet  near 
the  Gulf  of  Maine  (see  footnote,  p.  167),  for  the  tail 
of  the  latter  is  deeply  forked.     There  is  a  distinct 


Figure  79. — Silver  gar  (Tylosurus  marinus). 


168 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


longitudinal  ridge,  or  low  keel,  along  either  side  of 
the  caudal  peduncle.6 

Color. — Greenish  above,  silvery  on  the  sides, 
white  below;  a  bluish  silvery  stripe  aloDg  each  side 
becoming  broader  and  less  distinct  toward  the  tail; 
snout  dark  green;  there  is  a  blackish  blotch  deeper 
than  long  on  the  upper  part  of  the  cheek.  The 
fins  are  without  markings  for  the  most  part;  the 
dorsal  may  be  somewhat  dusky,  and  the  caudal 
bluish  at  its  base. 

Size. — The  silver  gar  grows  to  a  length  of  about 
4  feet. 

General  range. — Maine  to  Texas ;  abundant  along 
the  South  Atlantic  and  Gulf  coasts  of  the  United 
States,  often  running  up  fresh  rivers  above  tide 
water. 

Occurrence  in  the  Gulf  oj  Maine. — -The  silver  gar 
is  common  enough  along  the  southern  shores  of 
New  England,  e.  g.,  in  Rhode  Island  waters  and 
at  Woods  Hole  where  quite  a  few  are  found  from 
June  to  October.  Like  many  other  southern  fishes, 
however,  it  seldom  journeys  eastward  past  Cape 
Cod,  the  only  definite  records  of  it  within  the 
Gulf  of  Maine  being  of  several  collected  by  Dr. 
William  C.  Kendall  at  Monomoy  Island,  forming 
the  southern  elbow  of  Cape  Cod;  at  Wolfs  Neck, 
Freeport,  and  Casco  Bay,  Maine,  and  of  one 
found  by  Crane 6  in  the  stomach  of  a  tuna  that 
she  examined  at  Portland,  Maine,  in  July  1936. 
We  have  not  seen  it  in  the  Gulf,  nor  have  we  heard 
even  a  rumor  of  its  presence  there  from  fishermen, 
good  evidence  that  it  is  as  rare  a  straggler  as  the 
few  records  indicate,  for  large  silver  gars  are  not 


•  There  are  many  other  species  of  Ears  In  tropical  seas,  any  one  of  which 
might  stray  northward  with  the  Gulf  Stream  and  so  to  the  Gull  of  Maine. 
The  silver  gar  Is  Identifiable  among  them  by  the  following  combination  of 
characters  (no  one  character  alone  marks  It  out  among  Its  relatives):  mouth 
capable  of  being  nearly  closed;  caudal  peduncle  with  keels;  dorsal  and  anal 
fins  short,  the  former  with  13-17  rays,  the  latter  with  17-21  rays;  caudal  fin 
only  moderately  concave;  eyes  at  least  one-third  as  broad  as  the  head  Is  long 
behind  the  eyes;  body  not  excessively  slender  but  at  least  one- fifth  to  one-sixth 
as  deep  as  head  (including  Jaws)  Is  long;  body  not  very  strongly  compressed 
sldewlse;  Jordan  and  Evermann  (Bull.  47,  U.  8.  Nat.  Mus.,  Pt.  1,  1896.  p. 
709)  gives  a  useful  key  to  the  species  of  the  family. 

•  Zoologica,  New  York  Zool.  Soc,  vol.  21,  No.  16, 1936,  p.  211. 


fish  to  be  overlooked.  With  so  little  claim  to 
mention  here,  we  need  merely  note  that  it  is 
voracious,  feeding  on  all  sorts  of  smaller  fishes, 
and  that  it  runs  inshore,  possibly  even  into  river 
mouths,  to  spawn.  The  eggs,  described  by  Ryder,7 
are  about  3.6  mm.  (one-seventh  of  an  inch)  in 
diameter,  and  stick  together  and  to  any  object 
they  may  touch,  by  long  threads  scattered  over 
their  surface. 

Garfish  Ablennes  hiana  (Cuvier  and  Valenciennes) 
1846 

Jordan  and  Evermann,  1896-1900,  p.  718,  Athlennes 
hians.' 

Description. — -This  gar  resembles  the  silver  gar 
(Tylosurus)  closely  in  general  appearance,  as  well 
as  in  the  nature  and  arrangement  of  its  fins; 
especially  in  the  fact  that  the  rear  parts  of  its 
dorsal  and  caudal  fins  can  be  depressed  and  almost 
completely  concealed  in  a  groove,  with  the  for- 
ward parts  still  remaining  erect.  But  its  body  is  so 
strongly  flattened  sidewise  as  to  be  less  than  one- 
half  as  thick  as  it  is  deep,  instead  of  about  as 
thick  as  deep,  or  thicker,  as  it  is  in  the  silver  gar. 
The  dorsal  fin,  also,  arises  farther  back  relative  to 
the  anal  fin,  than  is  the  case  in  the  silver  gar. 
The  tail  fin  is  broadly  forked,  the  dorsal  and  anal 
fins  deeply  concave. 

Color. — -Back  greenish  with  bluish  green  reflec- 
tions; lower  part  of  sides  bright  silvery,  also 
the  abdomen;  snout  greenish;  dorsal  fin  mostly 
greenish,  but  with  the  rays  black-tipped;  tail  fin 
greenish.  Some  individuals  have  the  sides  plain 
silvery,  but  others  are  marked  with  dark  blotches 
or  indistinct  sooty  or  blue  crossbars.9 

Size. — Up  to  3  feet  long,  or  more. 


'  Bull.  U.  S.  Fish  Comm.,  vol.  1, 1882,  p.  283. 

«  The  original  spelling  was  Athlennes  (Jordan  and  Fordice,  Proa  U.  S.  Nat. 
Mus.,  vol.  9,  1886,  p.  342).  But  Jordan  and  Evermann  (Bull.  47,  U.  S.  Nat. 
Mus.,  Pt.  1,  1896,  p.  717,  footnote)  state  that  "Ablennes"  was  intended. 

•  Smith  (Sea  Fishes  of  Southern  Africa,  1949,  pi.  7,  fig.  26)  gives  a  colored 
Illustration  o  lone  with  blue  crossbars. 


^.iii  num  iuw.iiihhhu  wi.VMwm.1 1  iwp« 


Figure  80. — Garfish  (Ablennes  hians),  specimens  from  North  Truro,  Mass.,  and  from  Acapulco,  Mexico.     Drawing  by 

H.  B.  Bigelow. 


FISHES   OF  THE  GULF  OF   MAINE 


169 


General  range  and  occurrence  in  the  Gulf  of 
Maine. — Widespread  in  tropical  seas; 10  Brazil  to 
Chesapeake  Bay  in  the  western  Atlantic,  and 
northward  as  a  stray  to  Cape  Cod.    A  specimen 


of  this  tropical  fish,  about  23%  inches  (594  mm.) 
long  to  the  fork  of  the  tail,  was  taken  in  a  fish 
trap  on  the  shore  of  Cape  Cod  Bay  at  North 
Truro,  Mass.,  on  August  15,  1949.11 


THE  HALFBEAKS.     FAMILY  HEMIRAMPHIDAE 


The  halfbeaks  are  close  allies  of  the  billfishes 
(Belonidae,  p.  167),  but  it  is  only  the  lower  jaw 
that  is  greatly  prolonged  while  the  upper  jaw  is 
short  in  the  only  species  of  present  concern.  They 
are  largely  herbivorous,  feeding  mainly  on  green 
algae.  There  are  many  species  in  warm  seas, 
only  one  of  which  is  known  to  reach  the  Gulf  of 
Maine. 

Halfbeak  Hyporhamphus  unifasciatus 

(Kanzani)  1842 

Skipjack 

Jordan  and  Evermann,  1896-1900,  p.  721  (Hyporham- 
phus robeUi  (Cuvier  and  Valenciennes)  1846. 

Description. — The  most  striking  feature  of  this 
halfbeak  and  one  which  is  enough  of  itself  to  mark 
it  off  from  every  other  fish  known  from  the  Gulf 
of  Maine,  is  the  fact  that  while  the  lower  jaw  is 
very  long,  the  upper  jaw  is  short.12 

This  is  a  slender  fish,  its  body  only  one-sixth  to 
one-tenth  as  deep  as  long  (younger  fish  are  still 
more  slender),  tapering  slightly  toward  head  and 
tail.  Its  dorsal  (14  to  16  rays)  and  anal  (15  to 
17  rays)  fins  are  situated  far  back  and  opposite 
each  other,  as  in  the  silver  gar,  and  are  about 
equal  in  length  and  alike  in  outline.  There  are 
no  detached  finlets  between  them  and  the  caudal 

io  We  have  seen  specimens  from  Acapulco,  west  coast  of  Mexico;  Panama; 
Mauritius;  and  Zanzibar. 

11  This  specimen  was  presented  to  the  Museum  of  Comparative  Zoology  by 
John  Worthlngton  of  the  Pond  Village  Cold  Storage  Co.,  North  Truro, 
Mass. 

"  Should  a  halfbeak  be  taken  in  the  Gulf  of  Maine  it  would  be  well  to  con- 
sult Meek  and  Hildebrand  (Field  Museum  of  Natural  History,  Zool.  Series, 
vol.  15,  Pt.  1,  p.  232, 1923)  for  there  are  several  other  species  that  might  reach 
us  as  strays,  either  via  the  Gulf  Stream  route  or  from  offshore.  One,  indeed 
(Euleplorhamphus  veloz),  has  been  taken  at  Nantucket.  Its  lower  jaw  is  even 
longer  and  more  slender  than  that  of  the  halfbeak.  its  body  is  more  flattened 
sldewlse,  and  Its  pectoral  flns  are  longer. 


fin.  The  ventrals  stand  about  midway  between 
a  point  below  the  eye  and  the  base  of  the  caudal. 
The  teeth  are  small  and  the  scales  are  largest  on 
the  upper  surface  of  the  head.  The  beak  is  much 
shorter  in  young  fish  than  it  is  in  adults. 

Color. — Translucent  bottle  green  above  with 
silvery  tinge,  each  side  with  a  narrow  but  well- 
defined  silvery  band  running  from  the  pectoral  fin 
to  the  caudal  fin,  the  sides  darkest  above  and 
paler  below  this  band.  The  tip  of  the  lower  jaw  is 
crimson  in  life,  with  a  short  filament,  and  three 
narrow  dark  streaks  run  along  the  middle  of  the 
back.  The  forward  parts  of  the  dorsal  and  anal 
fins  and  the  tips  of  the  caudal  fins  are  dusky. 
The  lining  of  the  belly  is  black. 

Size. — Adults  are  seldom  more  than  1  foot  long. 

General  range. — Tropical  and  subtropical  on 
both  coasts  of  America  and  in  the  Gulf  of  Mexico ; 
abundant  off  the  South  Atlantic  United  States, 
not  uncommon  northward  to  Cape  Cod,  and  stray- 
ing to  the  coast  of  Maine. 

Occurrence  in  the  Gulf  of  Maine. — In  our  cool 
boreal  waters  the  halfbeak  is  only  a  rare  stray 
from  the  south,  previously  recorded  only  twice  in 
the  Gulf  of  Maine,  i.  e.,  from  Machias  and  from 
Casco  Bay,  Maine,  many  years  ago  in  each  case. 
We  can  now  add  one  taken  in  Quincy  Bay,  Boston 
Harbor,  July  10,  1951,  by  Gordon  Faust;  another 
off  Revere  (also  in  Boston  Harbor)  on  the  19th  of 
the  month,  by  John  M.  Hodson;  a  third,  taken 
in  a  trap  at  Sandwich,  September  24  of  the  same 
year,13  and  several  dozen  taken  in  a  pound  net  at 
Small  Point,  Maine,  July  14-15,  reportedjby  Leslie 
Scattergood. 


11  These  specimens  are  in  the  Harvard  Museum  of  Comparative  Zoology. 


Figure  81. — Halfbeak  (Hyporhamphus  unifasciatus),  Chesapeake  Bay. 

H.  L.  Todd. 


From  Jordan  and  Evermann.     Drawing  by 


170 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 

THE  NEEDLEFISHES.      FAMILY  SCOMBERESOCIDAE 


Both  jaws  (of  the  adult)  are  elongated  to  form  a 
slender  beak  in  the  needlefishes,  as  in  the  billfishes 
(p.  167),  and  the  anal,  dorsal,  and  ventral  fins  are 
set  far  back.  But  the  presence  of  several  finlets 
between  the  dorsal  and  anal  fins  and  the  caudal  in 
the  needlefishes  (which  the  billfishes  lack)  is  a 
ready  field  mark  for  their  identification.  Further- 
more, their  teeth  are  small  and  weak,  and  their 
bodies  only  moderately  slender.  Four  or  five 
species  are  known  in  warm  seas,  one  of  which  is  not 
uncommon  in  the  Gulf  of  Maine. 

Needlefish  Scomberesox  saurus  (Walbaum)  1792 

Billfish;  Skipper;  Saury 

Jordan  and  Evermann,  1896-1900,  p.  725. 

Description. — The  needlefish  resembles  the  sil- 
ver gars  in  its  slender  form  and  in  the  fact  tbat 
both  its  upper  and  lower  jaws  are  prolonged,  but 


greenish;  the  lower  parts  are  silvery  with  golden 
gloss.  Young  fry,  which  live  in  the  surface 
waters  of  the  open  Atlantic,  have  dark  blue  backs 
and  silvery  sides. 

Size. — Up  to  18  inches  long.  Those  caught 
along  Cape  Cod  run  a  foot  and  more  in  length. 

Habits. — The  skipper  is  an  oceanic  fish.  So 
far  as  known  it  always  lives  close  to  the  surface; 
so  much  so  indeed  that  in  English  waters,  where 
it  is  plentiful  in  summer,  few  are  caught  in  nets 
set  as  deep  as  a  fathom  or  two.  Its  hordes  are 
preyed  upon  by  porpoises  and  by  all  the  larger 
predaceous  fishes;  cod  and  pollock,  for  instance, 
feed  greedily  upon  them,  as  do  bluefish.  When 
they  strand  on  the  beaches,  as  often  happens,  it  is 
probably  while  they  are  fleeing  from  their  enemies. 
At  sea  they  attempt  to  escape  by  leaping,  whole 
companies  of  them  breaking  the  surface  together 
as  has  often  been  described,  and  as  we  have  seen 
them  doing  in  Massachusetts  Bay. 


Figure  82. —  Needlefish  (Scomberesox  saurus) .     Adult,  Woods  Hole.     From  Goode.     Drawing  by  H.  L.  Todd. 


it  differs  from  the  gars  in  having  a  series  of  five  or 
six  little  separate  finlets,  both  on  its  back  in  the 
space  between  the  dorsal  fin  and  the  caudal,  and 
on  its  lower  side  between  the  anal  fin  and  the 
caudal.  Its  body  is  about  nine  times  as  long 
(not  counting  caudal  fin)  as  deep,  flattened  side- 
wise,  tapering  toward  the  head  and  tail,  with 
slender  caudal  peduncle,  and  all  its  fins  are  small. 
Its  dorsal  fin  originates  slightly  behind  the  origin 
of  its  anal;  these  two  fins  are  alike  in  outline  and 
stand  far  back.  Its  ventrals  are  situated  about 
midway  the  length  of  the  body.  Its  caudal  is 
deeply  forked  and  symmetrical,  much  like  the 
tail  of  a  mackerel.  Its  trunk  is  covered  with 
small  scales  as  is  a  patch  on  each  gill  cover.  Its 
lower  jaw  projects  a  little  beyond  the  upper,  and 
its  teeth  are  pointed  but  very  small. 

Color. — Olive  green  above  with  a  silver  band 
on  each  side  at  the  level  of  the  eye  and  about  as 
broad  as  the  latter.  There  is  a  dark  green  spot 
above  the  base  of  each  pectoral;  the  dorsal  fin  is 


© 


Figure  83.- 


-Needlefish,  young,  about   2J4    inches  long. 
After  Murray  and  Hjort. 

It  is  not  likely  that  they  ever  spawn  in  the  cool 
waters  of  the  Gulf  of  Maine,  for  we  have  never 
taken  their  fry  in  our  tow  nets,  although  they  are 
among  the  most  numerous  of  young  fish  in  the 
open  Atlantic  between  the  latitudes  of  11°  or  12° 
N.  and  40°  N.  Although  their  eggs  are  covered 
with  filaments  like  those  of  the  silver  gars,14  they 
are  not  adhesive  as  the  latter  are,  but  drift  near 
the  surface.  The  most  interesting  phase  in  the 
development  of  the  skipper  is  that  its  jaws  do  not 
commence  to  elongate  until  the  fry  have  grown 

»  Skipper  eggs  were  so  described  by  Haeckel  (Archiv  fur  Anatomie,  Phy- 
siologic, and  Wissenschaftliche  Medecin,  1855,  p.  23,  pi.  5,  fig.  15,)  75  years 
ago.  They  were  not  seen  again  until  1910  when  similar  eggs,  2.2  mm.  In 
diameter,  covered  with  filaments,  were  towed  in  the  Atlantic  by  the  Michael 
Sart  (Murray  and  Hjort.  Depths  of  the  Ocean,  1912,  p.  742,  fig.  531). 


FISHES   OF  THE  GULF  OF   MAINE 


171 


to  about  1%  inches  (40  mm.),  and  that  the  lower 
jaw  out-strips  the  upper  at  first,  so  that  fry  of 
4  to  6  inches  (100  to  150  mm.)  look  more  like 
little  halfbeaks  ("Hemirampbus"  itage)  than  like 
their  own  parents. 

European  students  tell  us  that  the  skipper  feeds 
on  the  smaller  pelagic  Crustacea  and  probably 
also  on  small  fish,  for  it  is  sometimes  caught  on 
hook  and  line.  One  examined  by  Linton  at 
Woods  Hole  contained  chiefly  annelid  worms, 
fragments  of  fish,  copepods  and  crustacean  larvae, 
with  some  vegetable  debris. 

General  range. — Temperate  parts  of  the  Atlantic, 
Pacific,  and  Indian  Oceans,  known  in  the  open 
sea  as  far  north  as  northern  Norway  off  the  Euro- 
pean coast,  and  to  southern  Newfoundland  and 
southern  Nova  Scotia  16  off  the  eastern  American 
coast. 

Occurrence  in  the  Gulf  of  Maine. — -While  the 
skipper  is  a  straggler  to  our  Gulf  from  warmer 
waters  offshore  or  farther  south,  it  has  been 
taken  along  the  northern  coasts  of  New  England 
more  often  than  have  any  of  its  relatives ;  specific- 
ally along  Cape  Cod;  at  Provincetown ;  at  several 
locations  in  Massachusetts  Bay  where  we  have 
seen  schools  of  them;  at  Annisquam  a  few  miles 
north  of  Cape  Ann;  at  Old  Orchard  (Maine);  in 
Casco  Bay;  at  Monhegan  Island;  in  the  central 
part  of  the  Gulf; 16  among  the  islands  at  the 
northern  entrance  to  the  Bay  of  Fundy;  and  on 
the  northern  part  of  Georges  Bank,  where  one  was 
gaffed  from  the  Albatross  //on  September  20, 1928. 
But  we  find  no  record  of  it  along  the  Nova  Scotia 
shore  of  the  Gulf  of  Maine.     The  inner  curve  of 


Cape  Cod  from  Provincetown  to  Wellfleet  seems 
to  be  a  regular  center  of  abundance  for  it,  as 
Storer  long  ago  remarked,  for  schools  of  billfish 
are  picked  up  in  the  traps  along  that  stretch  of 
beach  almost  every  year,  the  catch  occasionally 
amounting  to  hundreds  of  barrels,  and  hosts  of 
them  have  been  known  to  strand  there.  Its  num- 
bers fluctuate  greatly  from  year  to  year,  however, 
and  it  often  fails  to  appear.17 

They  are  likely  to  be  taken  any  time  from  mid- 
June  to  October  or  November,  the  largest  catches 
usually  being  made  late  in  summer.18  We  have 
seen  several  schools  skipping,  as  is  their  common 
habit,  off  the  Scituate  shore  on  the  southern  side 
of  Massachusetts  Bay.  But  skippers  are  so  much 
less  common  farther  within  Massachusetts  Bay 
that  some  fishermen  had  never  heard  of  them 
there.  They  appear  only  as  strays  north  of  Cape 
Ann.  But  it  would  not  be  astonishing  if  a  large 
school  were  to  be  encountered  anywhere  within 
the  Gulf;  witness  their  occasional  abundance  off 
northern  Nova  Scotia.19  When  skippers  do  in- 
vade the  waters  of  our  Gulf,  they  may  be  expected 
in  multitudes,  for  they  usually  travel  in  vast 
schools.  Day,20  for  example,  mentions  the  cap- 
ture of  100,000  in  a  single  haul  in  British  waters. 

Commercial  importance. — The  skipper  is  not  of 
much  commercial  importance,  being  too  sporadic 
in  its  appearances.  However,  when  large  catches 
are  made  on  Cape  Cod  they  find  a  ready  sale  near 
by.  If  too  many  are  caught  for  the  local  trade  to 
absorb,  they  are  sent  to  Boston,  where  they  are 
sold  for  bait. 


THE  FLYING  FISHES.  FAMILY  EXOCOETIDAE 


The  typical  flying  fishes  have  one  dorsal  fin 
and  one  anal  fin,  both  of  them  soft  rayed,  both 
of  them  located  far  rearward,  and  with  the  anal 
below  the  dorsal.  Their  ventral  fins  are  well 
behind  their  pectorals,  their  tails  are  very  deeply 
forked  with  the  extreme  tips  rounded,  the  lower 
lobe  the  longer,  and  they  have  small  mouths  and 
large  rounded  scales.  Their  most  distinctive 
feature  is  that  their  pectorals  are  so  long  and  so 
stiff  that  their  owners  can  plane  through  the  air 

'» Cornish  (Contrib.  Canadian  Biol.,  (1902-1905)  1907,  p.  83)  states  that 
large  schools  can  often  be  seen  at  Canso,  Nova  Scotia,  skipping  over  the  water 
as  they  flee  from  the  pollock. 

'•  The  Museum  of  Comparative  Zoology  has  a  specimen,  taken  115  miles 
southeast  of  Portland  Lightship. 


on  them,  several  feet  above  the  water,  which  they 
do  mostly  in  attempts  to  escape  their  enemies, 
and  as  has  been  described,  time  and  again.  Jordan 
and  Evermann  have  given  a  popular  account  of 
this  so-called  "flight"  (really  not  flight  at  all, 
for  the  flying  fish  does  not  flap  its  wings)  in  their 


"  Blake  (American  Naturalist,  vol.  4,  1870,  p.  521)  remarked  that  while 
years  before  he  saw  thousands  stranded  at  Provincetown  not  one  was  seen 
in  1870.    It  failed  In  1921,  also,  and  no  doubt  in  many  intervening  years. 

"  We  are  Indebted  for  information  on  the  local  abundance  of  billfish  on 
Cape  Cod  to  Capt.  L.  B.  Ooodspeed,  a  fisherman  of  long  experience  and 
close  observation. 

a  Cornish  (Contrib.  Canadian  Biol.,  1902-1905  (1907),  p.  83)  states  that 
large  schools  can  often  be  seen  at  Canso  skipping  over  the  water  as  they  flee 
from  the  pollock. 

»  The  fishes  of  Great  Britain  and  Ireland,  vol.  2,  1880-1884,  p.  152. 


172 


FISHERY   BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


Guide  to  the  Study  of  Fishes,  1905,  p.  157.  For 
a  more  detailed  study,  based  similarly  on  first- 
hand observations,  we  refer  the  reader  to  Hubbs, 
Papers  of  the  Michigan  Academy  of  Sciences, 
vol.  17,  1933,  pp.  575-611.  Voyagers  in  tropical 
seas  are  perhaps  more  familiar  with  flying  fishes 
than  with  any  other  fishes.  And  they  are  often 
seen  in  the  warm  ultramarine-blue  waters  of  the 
Gulf  Stream  abreast  of  our  northeastern  coast. 
But  none  of  them  are  to  be  expected  in  the  boreal 
waters  of  our  Gulf  except  as  the  rarest  of  strays. 
A  flying  fish  could  hardly  be  mistaken  for  any- 
thing else,  except  possible  for  a  flying  gurnard 
(p.  472).  But  a  glance  should  be  enough  to  tell 
which  of  them  one  has  in  hand,  for  the  flying 
fishes  have  stiff,  narrow,  pointed  wings,  only  on 
dorsal  fin  and  a  very  deeply  forked  tail,  whereas 
the  so-called  wings  of  the  flying  gurnard  are  broad, 
rounded,  and  extremely  flexible;  they  have  two 
dorsal  fins,  and  a  tail  fin  that  is  only  weakly  con- 
cave in  outline. 

Flying  Fish  Cypselurus  heterurus(R&finesq\ie)  1810 
Jordan  and  Evermann,  1896-1900,  p.  2836. 

Distinctive  features  of  flying  fishes  of  the  genus 
Cypselurus,  among  its  tribe,  are  pectoral  fins  so 
long  that  they  overlap  the  anal  fin  considerably 
when  laid  back;  ventrals  standing  far  rearward 
and  nearly  or  quite  half  as  long  as  the  pectorals; 
anal  fin  with  its  point  of  origin  only  a  litle  forward 
of  the  mid-point  of  the  base  of  the  dorsal  fin;  and 
second  ray  of  the  pectoral  fins  branched.  The 
species  heterurus  has  a  plain  gray  dorsal  fin;  it  has 
no  teeth  on  the  palatine  bonej  in  the  roof  of  its 
mouth;  and  the  pale  edging  of  the  outer  margin 
of  its   pectoral   fins   is   extremely  narrow.     The 


number  of  its  scales  is  distinctive,  also,  as  is  the 
number  of  its  vertebrae.  But  these  last  two 
characters  are  matters  for  the  specialist. 

Color. — Dark  blue  gray  on  the  back  and  on  the 
upper  part  of  the  sides,  silvery  lower  down  on  the 
sides,  and  below;  the  dorsal  fin  is  plain  gray,  the 
rear  margin  of  the  pectorals  with  a  very  narrow 
pale  edging. 

Size. — The  species  heterurus  is  one  of  the  larger 
flying  fishes,  commonly  growing  to  a  length  of 
about  1  foot  (to  the  base  of  the  tail  fin). 

Occurrence  in  the  Gvlj  of  Maine. — A  flying  fish, 
about  9%  inches  long  to  the  fork  of  the  tail,  seem- 
ingly of  this  species  but  not  in  good  enough  con- 
dition for  certain  identification,  was  taken  in  a 
trap  of  the  Pond  Village  Cold  Storage  Co.  at 
North  Truro,  on  the  Massachusetts  Bay  shore  of 
Cape  Cod,  on  August  4,  1952.  This  is  thf  only 
record  of  one  of  its  tribe,  from  our  Gulf.  And 
the  only  record  of  a  flying  fish  from  Nova  Scotian 
coastal  waters  is  by  Jones,  of  one  taken  at  Sable 
Island,  in  1859. 

Flying  fishes  are  taken  now  and  then  at  Woods 
Hole,  the  species  heterurus  perhaps  more  often 
than  any  other,  according  to  published  report, 
but  several  of  the  kinds  to  be  expected  in  the  Gulf 
Stream  off  our  coast  resemble  one  another  very 
closely  indeed.  So  we  suggest  that  if  a  flying  fish 
should  be  taken  in  our  Gulf  that  does  not  seem  to  fit 
the  accompanying  illustration  (fig.  83A)  it  be 
forwarded  either  to  the  Fisheries  Laboratory  of  the 
U.  S.  Fish  and  Wildlife  Service,  Woods  Hole, 
Massachusetts;  to  the  Department  of  Fishes, 
U.  S.  National  Museum,  Washington,  D.  C;  or 
to  the  Department  of  Fishes,  Museum  of  Com- 
parative Zoology,  Cambridge,  Massachusetts,  to 
be  named. 


FiotTBE  83A. — Flyingfish  (Cypselurus  heterurus). 


After  Bruun  and  a  specimen  from  North  Truro,  Mass.     Drawing  by 
Jessie  Sawyer. 


FISHES   OF  THE  GULF  OF   MAINE 


173 


THE  SILVER  HAKE  AND  COD  FAMILIES— FAMILIES    MERLUCCIIDAE  AND  GADIDAE 


The  silver  hakes  and  the  cods  are  so  closely 
allied  that  many  European  ichthyoloigsts  group 
them  in  a  single  family.  American  practice,  how- 
ever, is  to  separate  them  because  of  certain  dif- 
ferences in  the  structure  of  the  skull  and  ribs. 
They  are  soft-finned  fishes,  lacking  true  spines  at 
any  stage  in  development  (though  in  one  local 
species,  the  silver  hake,  the  basal  parts  of  the 
dorsal  and  anal  fin  rays  are  so  stiff  as  to  feel  like 


spines  to  the  touch),  but  they  are  distinguishable 
from  all  other  soft-rayed  Gulf  of  Maine  fishes  by 
the  fact  that  their  large  ventral  fins  are  situated 
under  the  pectorals  or  in  front  of  them,  and  not 
behind  them,  as  they  are  in  the  herrings  and 
salmons.  They  and  their  relatives,  the  grenadiers 
(p.  243),  are  separated  from  most  of  the  typical 
spiny-rayed  fishes  by  the  structure  of  the  skull.21 


Key  To  Gulf  Of  Maine  Hakes,  Cods,  And  Other  Species 

1.  There  are  three  separate  dorsal  fins  and  two  anal  fins 2 

There  are  two  separate  and  well  developed  dorsal  fins 5 

There  is  only  one  well  developed  dorsal  fin 11 

2.  The  lateral  line  is  black;  there  is  a  black  blotch  on  each  shoulder -.- -Haddock,  p.  199 

The  lateral  line  is  pale;  there  is  no  shoulder  blotch 3 

3.  The  lower  jaw  projects  beyond  the  upper;  the  chin  barbel  is  very  small,  if  there  is  one Pollock,  p.  213 

The  upper  jaw  projects  beyond  the  lower;  the  chin  barbel  is  large 4 

4.  The  ventral  fins  are  narrow,  and  prolonged  as  filamentous  feelers  that  are  as  long  as  the  rest  of  the  fin;  the  eyes  are 

small..  Tomcod,  p.  196 

The  ventral  fins  are  broad,  and  their  filamentous  tips  are  less  than  one-third  as  long  as  the  remainder  of  the  fin;  the 
eyes  are  large Cod,  p.  182 

5.  The  anal  fin  originates  considerably  in  front  of  the  point  of  origin  of  the  second  dorsal  fin Hakeling,  p.  233 

The  anal  fin  originates  under  the  point  of  origin  of  the  second  dorsal  fin  or  behind  it 6 

6.  The  ventral  fins  are  short  and  of  ordinary  form Silver  hake,  p.  173 

The  ventral  fins  are  very  long  and  feeler-like 7 

7.  The  first  dorsal  fin  is  hardly  higher  than  the  second  dorsal,  and  none  of  its  rays  are  prolonged  or  filamentous 

Spotted  hake,  p.  230 

The  first  dorsal  fin  is  much  higher  than  the  second  dorsal,  with  one  or  two  long  filamentous  rays 8 

The  ventral  fins  reach  nearly  or  quite  as  far  back  as  the  rear  end  of  the  anal  fin Long-finned  hake,  p.  232 

The  ventral  fins  do  not  reach  back  to  the  middle  of  the  anal  fin 9 

The  anal  fin  is  so  deeply  notched  about  midway  of  its  length  as  to  suggest  two  separate  fins Blue  hake,  p.  233 

The  anal  fin  is  of  about  equal  height  from  end  to  end 10 

There  are  about  140  rows  of  scales  along  the  lateral  line  from  gill  opening  to  base  of  caudal  fin;  the  upper  jaw  bone 

reaches  back  to  below  the  rear  edge  of  the  eye White  hake,  p.  221 

There  are  only  about  110  rows  of  scales  along  the  lateral  line;  the  upper  jaw  bone  reaches  back  only  as  far  as  the 

rear  edge  of  the  pupil Squirrel  hake,  p.  223 

There  are  no  isolated  rays  in  front  of  the  dorsal  fin,  nor  barbels  on  the  top  of  the  snout Cusk,  p.  238 

The  dorsal  fin  is  preceded  by  a  fringe  of  short  rays  and  one  long  ray;  the  top  of  the  snout  bears  barbels  as  well  as  the 

chin 12 

There  are  three  barbels  on  the  top  of  the  nose Four-bearded  rockling,  p.  234 

There  are  only  two  barbels  on  the  top  of  the  nose ...Three-bearded  rockling,  p.  237 


8, 


10. 


11 


12. 


Silver  hake  Merluccius  bilinearis  (Mitchill)  1814 
Whiting;   New  England   hake 

Jordan  and  Evermann,  1896-1900,  p.  2530. 

Description. — The  presence  of  two  separate  and 
well  developed  dorsal  fins,  both  of  them  soft-rayed, 
the  second  much  longer  than  the  first,  combined 
with  the  location  of  the  ventrals  on  the  chest,  is 
sufficient  field  mark  to  distinguish  the  silver  hake 
from  all  other  Gulf  of  Maine  fishes  except  for  the 


true  hakes  (genus  Urophycis,  p.  221).  And  there 
is  no  danger  of  confusing  it  with  any  of  the  latter, 
for  it  lacks  the  chin  barbels  so  characteristic  of 
them,  and  its  ventrals  are  of  the  ordinary  finlike 
form,  whereas  those  of  the  true  hakes  are  altered 
into  long  feelers.  It  is  a  rather  slender  fish,  about 
five  to  six  times  as  long  as  it  is  deep,  its  body 
rounded  in  front  of  the  vent  but  flattened  sidewise 
behind  it,  with  large  flat-topped  head  occupying 


»>  The  hypercoracoid  bone  lacks  an  aperture  (technically  a  "foramen"). 


174 


FISHERY   BULLETIN    OF   THE    FISH   AND   WILDLIFE   SERVICE 


Figure  84. — Silver  hake  (Merluccius  bilinearis) .  A,  adult, 
Nova  Scotia,  from  Goode,  drawing-  by  H.  L.  Todd;  B, 
egg;  C,  larva,  6.5  mm.;  D,  larva,  11  mm.;  E.,  young  fry, 
23  mm.     B-E,  after  Kuntz  and  Radcliffe. 


about  one-fourth  of  the  total  length;  large  eyes; 
and  wide  mouth  armed  with  two  or  more  rows  of 
sharp  recurved  teeth,  and  with  the  lower  jaw 
projecting  beyond  the  upper. 

The  first  dorsal  fin  (11  to  14  rays)  originates 
close  behind  the  gill  openings,  is  roughly  an  equi- 
lateral triangle  in  shape,  and  is  separated  by  a 
short  space  from  the  second  dorsal.  The  second 
dorsal  (38  to  41  rays)  is  about  four  times  as  long 
as  the  first  dorsal,  but  hardly  more  than  half  as 
high,  and  is  of  distinctive  outline,  being  deeply 
emarginate  two-thirds  of  the  way  back,  with  the 
rear  section  the  higher  of  the  two.  The  anal  fin 
(38  to  41  rays)  corresponds  in  height  and  in  shape 
to  the  second  dorsal,  under  which  it  stands.  The 
caudal  fin  is  square  tipped  when  widespread,  but 
its  rear  margin  is  weakly  concave,  otherwise. 
The  pectorals  are  rather  narrow,  their  tips  slightly 
rounded,  and  they  reach  back  far  enough  to  over- 
lap the  second  dorsal  a  little.     The  ventral  fins, 


situated  slightly  in  front  of  the  pectorals,  are 
perceptibly  shorter  than  the  latter,  with  about 
half  as  many  rays  (7). 

Color. — The  silver  hake  is  dark  gray  above  of 
brownish  cast;  but  silvery-iridescent,  as  its  name 
implies,  or  with  golden  reflections.  The  lower 
part  of  its  sides  and  its  belly  are  silvery.  The 
inside  of  its  mouth  is  dusky,  the  lining  of  its  belly 
blackish.  The  fish  is  brightly  iridescent  when 
taken  from  the  water,  but  fades  soon  after  death. 

Size. — Maximum  size  about  2^  feet  long  and 
about  5  pounds  in  weight,  but  adults  average  only 
about  14  inches  long. 

Habits. — Silver  hake  are  strong  swift  swimmers, 
well  armed  and  extremely  voracious.  They  prey 
on  herring  and  on  any  other  of  the  smaller  school- 
ing fish,  such  as  young  mackerel,  menhaden,  ale- 
wives,  and  silversides.  Probably  a  complete  diet 
list  would  include  the  young  of  practically  all  the 
common  Gulf  of  Maine  fishes,  for  Vinal  Edwards 


FISHES   OF  THE  GULF  OF   MAINE 


175 


recorded  the  following  from  silver  hake  taken  at 
Woods  Hole:  alewife,  butterfish,  cunner,  herring, 
mackerel,  menhaden,  launce,  scup,  silversides, 
smelt,  also  the  young  of  its  own  species.  A  23^- 
inch  silver  hake,  taken  at  Orient,  N.  Y.,  had  75 
herring,  3  inches  long,  in  its  stomach.22  And  it 
is  probable  that  the  silver  hake  that  frequent 
Georges  Bank  feed  chiefly  on  young  haddock. 
They  eat  squid  when  occasion  offers.  The  small 
ones  in  particular  prey  regularly  on  large  shrimp 
(Pandalus)  in  the  deep  troughs  in  our  Gulf,  where 
experimental  trawlings  by  the  Atlantis  in  the 
summer  of  1936  took  about  four  times  as  many 
silver  hake  at  stations  where  these  shrimps  were 
abundant  as  at  stations  where  shrimp  were  scarce.23 
They  sometimes  take  crabs,  and  bite  freely  on 
almost  any  bait,  such  as  clams  or  cut  fish. 

Though  silver  hake  do  not  school  in  definite 
bodies,  multitudes  of  them  often  swim  together, 
and  such  bands  sometimes  drive  herring  ashore, 
and  strand  themselves,  in  the  pursuit.  Events  of 
this  sort  are  oftenest  reported  in  early  autumn 
when  the  spent  fish  are  feeding  ravenously  after 
the  effort  of  spawning,  but  this  may  also  happen 
at  any  time  during  the  summer.  Thus,  Prof.  A.  E. 
Gross  saw  the  beach  at  Sandy  Neck,  Barnstable, 
Mass.,  covered  with  them  on  several  occasions  in 
June  and  July  1920.24  Doctor  Huntsman  informs 
us  that  spent  fish  frequently  strand  on  the  beaches 
on  both  sides  of  the  Bay  of  Fundy  in  September. 
We  once  saw  an  army  of  silver  hake  harrying  a 
school  of  small  herring  on  a  shelving  beach  at 
Cohasset,  Mass.  We  half  filled  our  canoe  with 
pursuers  and  pursued,  with  our  bare  hands. 

It  is  said  that  European  silver  hake  rest  on  the 
bottom  by  day  and  hunt  by  night,  and  it  is  usually 
at  night  that  the  American  fish  run  up  into  the 
shallows  and  enter  the  traps.  But  strandings  also 
take  place  by  day.  Silver  hake,  like  many  other 
rapacious  species,  are  wanderers,  independent  of 
depth  within  wide  limits,  and  of  the  sea  floor. 
Sometimes  they  swim  close  to  the  bottom,  some- 
times in  the  upper  levels  of  the  water,  their  vertical 
movements  being  governed  chiefly  by  their 
pursuit  of  prey.  Their  upper  limit  is  the  tide  line; 
at  the  other  extreme  they  have  been  trawled 
repeatedly  as  deep  as  150  to  400  fathoms  on  the 
continental  slope  off  southern  New  England,  and 


as  deep  as  296  fathoms  off  North  Carolina.26 
When  they  are  on  bottom  they  are  caught  in- 
differently on  sandy  or  pebbly  ground,  or  on  mud 
(as  in  the  deep  trough  west  of  Jeffreys  Ledge, 
p.  175);  seldom  around  rocks. 

The  lowest  temperatures  in  which  we  have  known 
of  silver  hake  being  taken  have  been  between 
38°  and  40°  F.  (probably),  in  the  bottom  of  the 
deep  trough  west  of  Jeffreys  Ledge,  August  1936,2" 
about  40°  F.  (4.4°  C.)  at  28  fathoms  off  New 
York,  February  28,  1929,  and  about  39.5°  F. 
(4.2°  C.)  at  19  fathoms  in  the  same  general  region, 
February  5,  1930. w  And  most  of  the  winter  and 
early  spring  records  for  it  have  been  where  the 
bottom  temperature  was  warmer  than  about 
43°  F.  (6°  C.).28  At  the  other  extreme,  we  have 
never  heard  of  them  in  any  numbers  where  the 
water  was  warmer  than  about  64°  F.  (18°  C.) ;  the 
monthly  catches  made  in  Cape  Cod  Bay  (see  p. 
180)  are  especially  instructive  in  this  regard. 

Breeding  habits. — The  silver  hake  is  the  most 
important  summer  spawner  among  Gulf  of  Maine 
fishes  that  are  important  commercially,  just  as  the 
haddock  is  for  spring  and  the  pollock  for  autumn. 
The  Gulf  is  probably  its  most  prolific  nursery,  too, 
and  it  spawns  over  the  outer  part  of  the  Nova 
Scotia  Banks  also,  as  far  east  as  Sable  Island, 
Dannevig  29  having  recorded  large  egg  catches  in 
the  offing  of  Halifax.  But  this  is  probably  its 
eastern  breeding  limit,  for  the  Canadian  Fisheries 
Expedition  found  no  silver  hake  eggs  or  fry  on 
Banquereau  or  Misaine  Banks;  in  the  Laurentian 
Channel;  or  on  the  Newfoundland  Banks.  In  the 
opposite  direction,  eggs  in  fair  numbers  have  been 
taken  in  the  tow  nets  off  Woods  Hole  in  July  and 
August ;  the  Albatross  II  has  found  them  and  the 
resultant  larvae  near  shore  off  Long  Island  in 
June  and  July,  with  eggs  as  far  south  as  the  offing 
of  Cape  May ;  and  the  young  fry  have  been  caught 
off  New  York 30  from  spring  to  autumn. 

We  have  no  evidence  that  silver  hake  commence 
to  spawn  before  June,  north  of  Cape  Cod,  our 
earliest  egg  record  having  been  for  the  11th  of  that 


»  Nichols  and  Breder,  Zoologies,  N.  Y.  Zool.  Soc,  vol.  9, 1927,  p.  163. 
■  For  details,  see  Bigelow  and  Schroeder,  Biol.  Bull.,  vol.  76, 1939,  p.  315. 
*  The  Auk,  vol.  40, 1923,  p.  19. 


»  Goode  and  Bean,  Smithsonian  Contrib.  Knowl.,  vol.  30,  1895,  p.  387. 

*  No  temperature  was  taken,  but  38.6"  F.  (3.66°  C.)  was  recorded  there  at 
85  fathoms,  on  August  15, 1914,  and  39.8°  F.  (4.33°  C.)  at  72  fathoms  on  August 
15,  1913. 

•'  Specimens  trawled  by  Albatroti  II. 

"  AlbatroM  II  trawled  a  considerable  number  at  stations  scattered  along 
the  continental  slope,  from  the  offing  of  southern  New  England  to  the  offing 
of  Chesapeake  Bay,  in  February  1929  and  1930,  and  in  April  1930. 

»  Canad.  Fish.  Exped.  (1914-1915),  1919,  p.  27. 

"  Nichols  and  Breder,  Zoologies,  New  York  Zool.  Soc,  vol.  9, 1927,  p.  163. 


176 


FISHERY   BULLETIN   OF   THE    FISH   AND   WILDLIFE   SERVICE 


month ;  in  our  Gulf,  egg  production  (as  evidenced 
by  the  numbers  of  eggs  taken  in  our  tow  nets)  is  at 
its  height  in  July  and  August  and  continues 
through  September,  though  less  freely,  with 
October  22  as  our  latest  date.  Similarly,  the 
Canadian  Fisheries  Expedition  found  no  silver 
hake  eggs  in  Nova  Scotia  waters  east  of  Cape 
Sable  in  May,  but  many  in  July. 

It  is  impossible  to  establish  the  exact  tempera- 
ture at  which  silver  hake  are  spawning  at  any 
particular  station  without  knowing  at  what  level 
ripe  fish  are  in  the  water,  which  may  be  anywhere 
between  the  surface  and  the  bottom  with  this 
species.  It  may  be  definitely  stated,  however, 
that  they  never  spawn  in  as  cold  water  as  cod  and 
haddock  usually  do  in  the  western  Atlantic.  In 
1915,  for  example  (a  representative  season),  it 
was  not  until  the  entire  column  of  water  was 
slightly  warmer  than  41°  F.  at  the  locality  in 
question  that  we  found  the  first  silver  hake  eggs 
in  our  Gulf.  And  if  the  parent  fish  were  in  the 
upper  water  layers,  as  they  may  have  been,  all  the 
rich  spawnings  we  encountered  in  the  Gulf  during 
that  year,  and  during  the  next,  took  place  in  tem- 
peratures considerably  higher  still.  Similarly,  the 
silver  hake  eggs  towed  off  Halifax  by  the  Canadian 
Fisheries  Expedition  in  July  1915,  and  off  Shel- 
burne,  Nova  Scotia,  by  the  Grampus  on  September 
6  of  that  same  year  may  have  been  spawned  in 
water  warmer  than  50°  F.,  there  being  no  need  to 
assume  that  the  parent  fish  were  lying  in  the 
colder  bottom  stratum.  As  the  spawning  season 
draws  to  its  close,  in  September  and  October,  the 
minimum  temperatures  for  most  of  our  egg  stations 
have  been  higher  than  46°,  with  one  (our  latest 
record  for  the  season)  as  warm  as  57°  F.  at  all 
depths.  These  data  point  to  41°  to  45°  F.  as  the 
lowest  temperature  limit  for  the  spawning  of  the 
silver  hake,  with  most  of  the  eggs  produced  at 
45°  to  55°  F. 

In  the  case  of  any  fish  producing  buoyant  eggs 
the  tendency  of  the  latter  to  rise  (unless  counter- 
acted by  active  vertical  circulation  of  the  water) 
insures  that  their  development  shall  take  place  at 
the  temperature  of  the  upper  stratum  of  water, 
not  at  that  of  the  deeper  levels  where  they  were 
spawned.  And  the  silver  hake  is  no  exception  to 
this  rule.  While  we  have  towed  its  eggs  in  June, 
when  the  surface  was  still  only  about  42°  F.,  most 
of  the  egg  records,  and  all  our  rich  catches,  were 
all  made  where  the  upper  5  fathoms  or  so  were 


warmer  than  50°  and  usually  warmer  than  55°  F., 
with  the  temperature  of  the  immediate  surface 
60°  or  higher  in  most  cases.  Similarly,  silver 
hake  eggs  taken  off  Halifax  by  the  Canadian 
Fisheries  Expedition  in  July  1915,  and  off  Shel- 
burne,  Nova  Scotia,  by  the  Grampus  on  September 
6  of  that  year,  may  well  have  been  in  water  at 
least  as  warm  as  53°  F.,  there  being  no  reason  to 
suppose  they  were  far  below  the  surface.3'  All 
this  suggests  that  incubation  does  not  proceed 
normally  in  water  cooler  than  about  50°,  and  that 
it  is  most  successful  in  temperatures  as  high  as 
55°  to  60°  F.  This  evidence  that  while  the  eggs 
of  the  silver  hake  may  be  spawned  in  low  tem- 
peratures, a  comparatively  warm  surface  layer  is 
necessary  for  their  later  development,  offers  a 
reasonable  explanation  for  the  failure  of  this  fish 
to  breed  successfully  along  the  New  Brunswick 
shore  of  the  Bay  of  Fundy,  where  active  vertical 
circulation  maintains  surface  temperatures  as  low 
as  50°  to  55°  F.  throughout  the  summer,  at  least 
in  most  years.  At  the  other  extreme,  the  failure 
of  the  eggs  that  had  been  fertilized  artificially  to 
develop  in  the  hatchery  at  Woods  Hole  in  August 
temperatures  points  to  65°  to  70°  F.  as  the  upper 
limit  to  successful  incubation. 

According  to  Kuntz  and  Kadcliffe 32  only  part 
of  the  eggs  mature  at  one  time,  but  we  know  of 
no  estimate  of  the  number  of  eggs  a  single  female 
may  produce.  The  eggs  are  buoyant,  transparent, 
about  0.88  to  0.95  mm.  in  diameter,  with  a  single 
yellowish  or  brownish  oil  globule  of  0.19  to  0.25 
mm.  Incubation  is  rapid;  Kuntz  and  Radcliffe 
assumed  a  duration  of  48  hours  at  Woods  Hole, 
but  it  has  not  been  determined  for  the  cooler 
waters  in  the  Gulf  of  Maine.  The  larvae  are 
about  2.8  mm.  long  at  hatching,  slender,  with 
small  yolk  sac,  and  they  are  made  recognizable 
by  the  fact  that  the  vent  is  located  on  one  side, 
near  the  base  of  the  larval  fin  fold,  as  is  the  general 
rule  in  the  cod  family,  not  at  its  margin  as  in  most 
larval  fishes,  and  that  the  trunk  behind  the  vent 
is  marked  with  two  black  and  yellow  cross  bars. 
The  dorsal  and  anal  fins  and  the  caudal  fin  have  all 
assumed  their  definite  outlines  by  the  time  the 
little  fish  is  10  to  11  mm.  long,  and  fry  of  20  to 
25  mm.  begin  to  resemble  their  parents  in  general 
appearance. 

»  These  catches  were  all  made  either  at  the  surface  or  in  oblique  hauls 
with  open  nets. 

»  Kuntz  and  Radclifle  (Bull.  V.  S.  Bur.  Fish.,  vol.  35, 1918,  p.  109)  describe 
the  spawning  and  early  development. 


FISHES   OF  THE  GULF   OF   MAINE 


177 


Evidently  the  young  silver  hake  take  to  the 
deeper  water  layers  toward  the  end  of  their  first 
summer  or  that  autumn,  when  about  1  to  3  inches 
long,  for  fry  as  small  as  this  have  been  trawled  in 
good  numbers  off  southern  New  England  at  150 
fathoms  and  deeper  at  that  season  during  the  early 
explorations  of  the  U.  S.  Fish  Commission,33  by 
the  Albatross  II  off  Rockaway  and  off  Long  Beach, 
N.  Y.,  in  November  1928.  By  February  they 
may  be  anywhere  from  1%  to  5  inches  long,  de- 
pending on  how  early  they  are  hatched,  on  when 
they  take  to  bottom,  and  on  the  feeding  conditions 
they  find  there.34  They  may  be  anywhere  from 
about  2  inches  long  to  aboit  6%  inches  long  by 
April.36 

The  sizes  of  the  many  small  silver  hake  that  we 
have  collected  at  different  times  of  the  year,  both 
within  the  Gulf  of  Maine  and  southward  as  far  as 
the  offing  of  Chesapeake  Bay,  suggest  that  they 
reach  an  average  length  of  5%  to  7%  inches  when 
1  year  old,  and  of  about  9  %  to  11  inches  at  2  years 
of  age,  i.  e.,  in  their  third  summer.38  Fish  of  11 
to  14  inches  that  dominated  the  pound-net  catches 
made  near  PrOvincetown,  August  1939,37  were 
three-year-olds,  probably.  The  rate  of  growth  has 
not  been  traced  for  the  older  fish,  nor  can  it  be 
deduced  from  that  of  the  European  silver  hake  for 
the  latter  grows  to  a  considerably  greater  length, 
averaging  as  much  as  30  inches  at  8  years  in  the 
extreme  northerly  part  of  its  range  (Iceland)  and 
considerably  larger  still,  in  the  southern  part  (Gulf 
of  Gascony  and  off  Morocco).38  But  it  is  reason- 
able to  assume  that  the  growth  of  the  American 
fish  varies  similarly  with  the  latitude  (i.  e.,  that  it 
is  most  rapid  in  high  temperatures)  and  that  the 
American  female,  like  the  European,  grows  faster 
than  the  male.  The  European  Merluccius  ma- 
tures at  2  years,  which  is  probably  true  of  the 
American  species  as  well. 

General  range. — Continental  shelf  of  eastern 
North  America,  northward  to  the  Newfoundland 


»  Qoode,  Fish,  and  Fish.  Ind.  U.  8.,  Sect.  1, 1884,  p.  242. 

*•  Fry  taken  In  February  of  different  years  by  Albatross  II  have  ranged 
from  lii  inches  (31  mm.)  long  to  i%  inches  (120  mm.). 

«  In  April  1930  Albatross  II  trawled  young  fry  ranging  in  length  from  2 
Inches  (54  mm.)  to  6%  inches  (163  mm.)  long  at  a  number  of  stations  from 
the  offing  of  Rhode  Island  to  the  offing  of  Chesapeake  Bay,  at  14  to  85  fathoms. 

»  For  further  details,  see  Bigelow  and  Schroeder  (Biol.  Bull.,  vol.  76, 1939, 
pp.  319-320,  flg.  8). 

"  Information  supplied  by  Wm.  A.  Ellison,  Jr. 

"  Belloc,  Notes  et  Memolres  No.  21,  Office  Scientlflque  et  Technique  des 
Pfiches  Maritimes,  France,  1923. 


Banks,  southward  to  the  offing  of  South  Carolina;39 
most  abundant  between  Cape  Sable  and  New 
York.  It  is  represented  farther  offshore  and  in 
the  Gulf  of  Mexico  by  forms,  the  relationship  of 
which  to  the  Merluccius  of  our  northeastern  coast 
has  not  yet  been  determined.  The  silver  hake  is 
represented  in  Europe  by  a  close  relative,  the 
European  hake  (Merluccius  merluccius),  an  excel- 
lent account  of  the  natural  history  and  migrations 
of  which  is  given  by  Le  Danois.40 

Occurrence  in  the  Gulf  of  Maine. — Silver  hake 
are  familiar  fish  all  around  the  coasts  of  the  Gulf 
of  Maine  from  Cape  Cod  to  the  Bay  of  Fundy  and 
to  the  west  coast  of  Nova  Scotia.  But  it  has  long 
been  a  matter  of  common  knowledge  that  their 
chief  center  of  abundance  is  in  the  southwestern 
part  of  the  Gulf.  Thus  in  1945  (most  recent  year 
for  which  detailed  regional  statistics  are  available), 
the  reported  landings  were  between  46  and  47 
million  pounds  41  from  off  eastern  Massachusetts 
in  general,  including  the  shores  of  Cape  Cod  out  to 
the  western  slope  of  the  so-called  South  Channel, 
contrasting  with  only  about  4  million  pounds  for 
the  western  and  central  coasts  of  Maine,  and  with 
only  about  6,500  pounds  for  eastern  Maine. 
Silver  hake,  it  is  true,  are  said  to  be  common  in 
the  Passamaquoddy  region  (more  so  in  some  years 
than  in  others),  also  around  Grand  Manan  at  the 
mouth  of  the  Bay  of  Fundy.  And  they  are  re- 
ported at  various  localities  along  the  Nova  Scotia 
side  of  the  Bay  and  along  western  Nova  Scotia. 
But  they  are  not  mentioned  in  the  statistics  of  the 
Canadian  catches  for  these  waters,  hence  cannot 
be  very  plentiful  there. 

Silver  hake  are  numerous  over  the  west-central 
deeps  of  the  Gulf  also;  in  fact  we  found  this  the 
most  plentiful  fish  at  70  to  90  fathoms  in  the 
basins  off  Cape  Cod  in  the  southwestern  part  of 
the  Gulf  and  off  Mount  Desert  in  the  northeastern, 
in  August  1936;  also  in  the  trough  west  of  Jeffreys 
Ledge,  where  the  catches  of  them  averaged  292 
fish  (maximum  840,  niinimum  1)  as  reduced  to  the 
common  standard  of  one  hour's  trawling  with  an 
82-foot  shrimp  trawl.  And  it  is  interesting  that 
the  catch  there  averaged  about  four  times  as  great 


'•  The  silver  hake  has  been  said,  repeatedly,  to  range  southward  to  the 
Bahamas,  in  deep  water,  following  Jordan  and  Evermann  (Bull.  47,  U.  S. 
Nat.  Mas.,  Ft. 3, 1898,  p.  2530).  But  the  most  southerly  positive  record  we 
have  found  for  it  is  off  Charleston,  S.  C.  (Blake  Sta.  313,  lat.  32°  32'  N.,  long. 
78°  45'  W.;  Goode  and  Bean,  Smithsonian  Contrib.  Knowl.,  vol.  30,  1895, 
p.  387). 

«  Notes  et  Mem.,  2,  Off.  Sci.  Tech.  PGches  Maritimes,  France,  1920. 

«'  "Round"  and  dressed  fish  combined. 


178 


FISHERY   BULLETIN    OF   THE    FISH    AND   WILDLIFE   SERVICE 


(494  fish)  at  the  stations  where  shrimp  (Pandalus) 
were  plentiful  as  at  the  stations  where  these  were 
scarce  (114  fish),  evidence  that  silver  hake  con- 
gregate where  feeding  conditions  are  good.42 

Reported  landings  throw  little  light  on  the 
numbers  of  silver  hake  that  frequent  the  offshore 
rim  of  our  Gulf,  both  because  the  otter  trawls 
used  there  are  so  large-meshed  that  many  pass 
through,  and  because  most  of  those  that  are 
caught  on  Georges  and  Browns  Banks  are  thrown 
overboard  when  the  price  is  low.43  Experimental 
trawling,  however,  on  Georges  Bank,  April  to 
September  1913,  yielded  about  one-seventh  as 
many  silver  hake  on  the  average  (about  1,800 
fish)  as  haddock  (about  14,000  fish)  per  trip,  and 
the  Albatross  III  caught  an  average  of  about  150 
silver  hake,  running  about  one-half  pound  in 
weight,  per  trawl  haul,  in  250  hauls  on  various 
parts  of  Georges  Bank,  July,  August,  and  Septem- 
ber of  1948,  1949,  and  1950.  Thus  they  are 
moderately  plentiful  at  least  over  Georges  Bank 
as  a  whole,  and  there  is  no  reason  to  doubt  that 
this  applies  to  Browns  Bank  equally. 

These  catches  do  not  suggest  any  definite  con- 
centration on  any  one  part  of  the  bank,  at 
least  for  summer,  except  that  the  largest  that 
were  made  on  its  northern  part  were  in  hauls  from 
shoaler  than  30  fathoms,  whereas  the  largest 
catches  on  the  southern  part  were  in  hauls  from 
deeper  than  60  fathoms,  a  difference  which  may 
well  have  been  a  matter  of  the  food  supply.44  In 
April,  however,  of  1950,  the  silver  hake  were  not 
only  more  plentiful  along  the  northern  edge  of  the 
bank  (average  305  per  haul)  than  on  the  southern 
part  (average  77  per  haul)  but  so  strictly  confined 
to  the  deeper  levels  that  the  total  yield  of  66  trawl 
hauls  at  shoaler  than  60  fathoms  was  only  1 1  fish, 
contrasting  with  an  average  catch  of  232  fish  per 
haul  at  60  fathoms  and  deeper  (25  hauls).46 

Silver  hake  spawn  along  the  entire  coastal  zone 
from  Cape  Cod  to  Grand  Manan,  as  proved  by  the 
locations  of  the  egg  catches  (fig.  85).     The  sloping 


«  For  further  details,  see  Bigelow  and  Schroeder  (Biol.  Bull.,  vol.  76,  1939, 
p.  308,  table  1;  p.  314,  table  6. 

*>  Reported  landings,  1945-1947,  ranged  between  3,000  and  about  33,000 
pounds  for  Georges  Bank,  between  0  and  6,000  pounds  for  Browns. 

«  The  average  catch  per  haul  was  262  fish  from  shoaler  than  30  fathoms  and 
161  flsh  from  deeper  than  60  fathoms  on  the  northern  part  of  the  bank;  90  fish 
per  haul  from  shoaler  than  30  fathoms  and  286  flsh  per  haul  from  deeper  than 
60  fathoms  on  the  southern  part. 

<*  Twenty-one  trawl  hauls  at  60  fathoms  and  shoaler  yielded  none  at  all  in 
March;  but  no  hauls  were  made  in  that  month  deeper  than  60  fathoms,  where 
the  silver  hake  doubtless  were. 


sandy  bottom  around  the  northern  extremity  of 
Cape  Cod  and  off  the  eastern  slope  of  the  Cape 
evidently  is  an  important  center  of  reproduction. 
Thus  we  found  an  abundance  of  eggs  off  Race 
Point  on  July  7,  1915;  our  tow  nets  yielded  many 
eggs  at  two  stations  off  the  outer  shore  of  the  Cape 
on  July  22  of  the  following  year,  when  a  15-minute 
tow  there  at  20  fathoms,  with  a  net  one  meter  in 
diameter,  produced  approximately  25,000  larvae 
of  3  to  7  mm.,  the  richest  haul  of  young  fish  we 
have  ever  made  in  our  Gulf.  And  the  fish  were 
still  spawning  there  a  month  later,  as  proved  by  the 
presence  of  eggs. 

Other  occasions  when  we  have  taken  silver-hake 
eggs  in  large  numbers  have  been  off  Duck  Island 
near  Mount  Desert  on  July  19  and  on  August  18, 
1915;  near  Monhegan  Island,  August  4,  1915;  off 
Wooden  Ball  Island  near  the  mouth  of  Penobscot 
Bay  on  August  6,  1915;  and  off  Rye,  N.  H.,  on 
Jul}7  23  of  that  same  year.  But  we  have  never 
found  them  in  any  number  in  Massachusetts  Bay 
though  some  eggs  have  been  taken  there  on  several 
occasions  (fig.  85). 

Unfortunately,  no  quantitative  hauls  were  made 
at  any  of  the  more  productive  egg  stations,  hence 
the  number  of  silver-hake  eggs  present  in  the 
water  cannot  be  approximated.  But  the  vertical 
net  yielded  about  190  eggs  per  square  meter  of 
sea  surface  at  one  station  in  the  eastern  basin. 

Apparently  the  silver  hake  does  not  breed  suc- 
cessfully in  the  northern  side  of  the  Bay  of  Fundy 
for  neither  its  eggs  nor  its  fry  have  ever  been 
found  there.  But  the  capture  of  a  few  eggs  in 
Petit  Passage  in  our  tow  nets  on  June  10,  1915, 
suggests  that  it  may  spawn  on  the  southern  side 
of  the  bay  as  the  cunner  does  (p.  478).  And  it  may 
be  expected  to  do  so  along  the  west  coast  of  Nova 
Scotia,  for  the  Canadian  Fisheries  Expedition 
found  eggs  at  several  stations  off  outer  Nova 
Scotia,  eastward  to  the  longitude  of  Canso. 

The  presence  of  silver  hake  on  Georges  Bank 
throughout  the  summer  is  presumptive  evidence 
of  local  spawning,  though  we  have  taken  no  silver 
hake  eggs  or  larvae  there. 

The  locations  where  we  have  found  its  eggs 
suggest  that  the  silver  hake,  in  the  Gulf  of  Maine, 
spawns  chiefly  in  water  shoaler  than  50  fathoms. 
But  we  have  made  one  rich  haul  of  its  eggs  in  the 
center  of  the  eastern  basin.  And  the  discovery  of 
its  eggs  over  the  continental  slope  off  Nova  Scotia 


FISHES   OF  THE  GULF  OF   MAINE 


179 


*&T  °° 


/  .• 

/  : 

/  : 

/    .• 


70°  60° 

Figure  85. — Localities  where  eggs  (#),  and  larvae  (O)  of  silver  hake,  or  both  (O)  have  been  taken. 


by  the  Canadian  Fisheries  Expedition,46  with  the 
presence  of  ripe  fish  as  well  as  of  spent,  in  depths  as 
great  as  150  fathoms  and  more  off  southern  New 
England  47  proves  that  it  spawns  over  deep  water 
also.  The  European  silver  hake  usually  spawns 
in  50  to  100  fathoms. 

All  our  records  for  the  free-drifting  larvae  of 
the  silver  hake  in  the  Gulf,  unlike  those  for  its 
eggs,  have  been  in  the  southwestern  part.  And  we 
have  towed  along  the  coast  of  Maine  so  often  in 
August,  September,  and  October  (when  the  larvae 
spawned  from  June  to  August  might  have  been 
expected)  that  our  failure  to  find  them  east  of 
Cape  Elizabeth  seems  sufficient  evidence  that 
they  actually  are  limited,  in  their  regular  occur- 
rence, to  the  southwestern  part  of  the  Gulf 
(they  parallel  the  pelagic  stages  of  the  cod,  the 
haddock,  and  the  flatfishes  in  this)  and  to  the 
waters  westward  from  Cape  Cod.  Dannevig, 
too,  has  called  attention  to  the  absence  of  larvae 
of  the  silver  hake  in  Nova  Scotia  waters,  con- 


trasted with  the  presence  of  their  eggs  there.48 
One  possible  explanation  for  this  contrast 
between  larvae  and  eggs  is  that  it  may  mirror  the 
relative  percentage  of  eggs  that  hatch  in  the 
regions  in  question.  A  more  likely  explanation 
we  think,  when  taken  with  other  similar  facts 
of  distribution,  is  that  it  results  from  a  peripheral 
drift  around  the  shores  of  the  Gulf  from  north- 
east to  southwest,  in  which  the  eggs  take  part 
first  and  then  the  resultant  larvae.  This  type  of 
circulation,  in  fact,  has  been  established  so  well 
for  our  Gulf  by  hydrographic  evidence,  that  some 
such  involuntary  migration  is  inevitable,  not  only 
for  various  buoyant  fish  eggs  and  larvae  that  are 
produced  near  the  coast  line,  but  likewise  for  the 
drifting  communities  of  invertebrates,  and  of 
plants. 

It  is  now  known  that  large  numbers  of  the  silver 
hake  that  descend  to  the  deeper  water  layers  in 
the  southwestern  part  of  the  Gulf  during  their 
first  autumn  remain  there  during  the  following 


"  Dannevig,  Canadian  Fish.  Exped.,  (1914-15)  1919,  p.  28. 
1  Ooode,  Fish.  Ind.  U.  8.,  Sect.  1, 1884,  p.  242. 


<•  Canad.  Fish.  Eiped.  (1914-1916)  1919,  p.  28. 


180 


FISHERY  BULLETIN   OF  THE    FISH   AND   WILDLIFE   SERVICE 


year,  some  of  them  still  longer.  Thus  our  ex- 
perimental trawlings,  in  August  1936,  yielded 
large  numbers  of  the  1-year-olds  at  70  to  90 
fathoms  in  the  deep  basins  off  Cape  Cod  and 
west  of  Jefferys  Ledge;  also  off  Mount  Desert.4' 
And  nearly  all  of  the  silver  hake  that  come  close 
enough  inshore  to  enter  the  traps  in  the  south- 
western part  of  the  Gulf,  or  to  strand  on  the 
beaches  there,  are  good  sized  individuals  of  9 
inches  aDd  larger.  In  fact,  the  only  instance 
that  has  come  to  our  notice  of  any  considerable 
inshore  catch  of  one-year-old  fish  (about  6  to  8 
inches  long)  in  the  Cape  Cod  Bay  region  was 
near  Provincetown,  August  1939,  when  about 
1,900  of  them  were  taken  during  a  14-day  period.60 
Huntsman,  however,  reports  all  sizes  near  shore 
from  yearlings  on,  in  the  Passamaquoddy  region 
to  the  northeast,  and  in  the  neighboring  parts 
of  the  Bay  of  Fundy. 

The  silver  hake  3  years  old  and  older,  that 
provide  the  commercial  catches,  sometimes  ap- 
pear in  the  Cape  Cod  Bay-northern  Massachusetts 
region  as  early  as  the  last  week  in  March,  regularly 
by  May.  Welsh  saw  some  fish,  for  example,  in 
Ipswich  Bay  in  March  and  April  in  1913  (a  fairly 
representative  season),  considerable  numbers  in 
May,  aod  an  abundance  in  June.  And  this  may 
be  taken  as  typical  for  the  whole  coast  line  of 
the  Gulf  south  of  Portland;  also  for  Georges 
Bank,  where  the  first  silver  hake  were  taken  by 
the  otter  trawlers  from  April  27  to  29  in  1913, 
and  on  almost  every  trip  thereafter.  We  have 
not  beeD  able  to  learn  how  early  silver  hake  ap- 
pear on  the  coast  of  Maine  east  of  Portland, 
or  off  western  Nova  Scotia,  where  it  is  only 
within  the  past  few  years  that  any  attention  has 
been    paid   them. 

Around  Cape  Cod  Bay,  silver  hake  are  usually 
the  most  plentiful  in  June ;  disappear  more  or  less 
during  August  and  September;  and  reappear  in 
numbers  in  October,  though  far  fewer  then  than 
in  June,  as  is  illustrated  by  the  average  monthly 
catches  made  by  a  set  of  eight  traps  at  North 
Truro,  for  the  years  1946-1947  and  1950: 51  June, 
185,200  pounds;  July,  36,700  pounds;  August, 
1,206  pounds;  September,  1,780  pounds;  and 
October,  10,852  pounds. 

"  For  further  details,  see  Blgelow  and  Schroeder,  Biol.  Bull.  vol.  76,  1939, 
pp.  308,  319-320,  fig.  8. 

"  Information  supplied  by  William  A.  Ellison,  Jr. 

"Information  supplied  by  the  Pond  Village  Cold  Storage  Co.,  North 
Truro,  Mass. 


Whether  their  withdrawal  thence  in  summer  is  a 
matter  of  food,  or  whether  they  move  deeper  to 
escape  the  heat  of  summer  is  a  question  for  the 
future.  Farther  offshore  in  the  western  side  of 
the  Gulf,  and  to  the  northward,  silver  hake  are 
about  as  plentiful  in  July  and  August  as  they  are 
in  June,  as  indicated  by  the  vessel  landings  at 
Boston  and  Gloucester;  somewhat  less  so  in  Sep- 
tember and  October.  And  what  little  information 
we  have  suggests  that  summer  is  the  season  of 
greatest  plenty  for  them  in  the  Bay  of  Fimdy 
region,  though  there  are  far  fewer  of  them  there. 

The  great  majority  of  the  silver  hake  vanish 
from  the  inshore  waters  of  the  Gulf  during  the  late 
autumn,  November  seeing  the  last  of  them  in 
Massachusetts  and  Cape  Cod  Bays,  according  both 
to  our  own  observations  and  to  general  report. 
The  latest  catches  made  on  Georges  Bank  during 
the  experimental  trawlings  of  1913  were  on  De- 
cember 3  and  12.  And  though  a  few  are  brought 
in  from  the  grounds  off  Massachusetts  and  Cape 
Cod  during  January,  February,  and  March,  the 
catches  average  less  than  K70  as  great  for  those 
months  as  for  the  period  May  through  October, 
as  illustrated  by  the  monthly  landings  by  trawlers 
at  Boston  and  Gloucester  for  1947: M 


1,400 
2,  255 
1,  700 
7,  540 

May.. _       860,000 

June 1,  158,000 


January.. 
February. 

March 

April 


July 4,444,000 

August 4,  879,  000 

September 1,974,000 

October.. 2,381,000 

November 438,  000 

December 207,  000 

It  is  probable  that  the  fish  of  the  year  and  those 
that  are  only  1  year  old  winter  in  the  deeper  de- 
pressions near  where  they  first  took  to  the  bottom. 
It  is  unlikely  that  fish  as  small  fish  as  those  we 
have  trawled  in  these  situations,  in  August,  can 
travel  far. 

The  wintering  ground  of  the  Gulf  of  Maine  pop- 
ulation of  larger  silver  hake  is  not  known.  Many  of 
them  may  winter  near  the  sea  floor  in  the  deep  open 
troughs  of  the  Gulf,63  where  the  bottom  water  at 
75  to  100  fathoms  and  deeper  continues  warmer 
than  39°  F.  (4°  C.)  even  at  the  coldest  time  of 
year.  Evidence  in  this  direction  is  that  it  is  only 
deeper  than  60  fathoms  that  good  April  catches 
have  been  reported  on  Georges  Bank  (p.  180).  It 
is  also  possible  that  part  of  them  move  out  to 
the  shelf  off  southern  New  England  to  winter,  or 


u  Pounds  of  round  fish  and  dressed  fish  combined. 

"  Practically  no  trawling  is  done  m  winter  in  the  deepest  parts  of  the  Gulf. 


FISHES   OF  THE  GULF  OF   MAINE 


181 


even  to  the  continental  slope  as  the  European 
silver  hake  do.  Scattered  catches,  in  fact,  of  half- 
grown  silver  hake  and  larger  are  made  by  otter 
trawlers  off  southern  New  England,  and  off  New 
York  in  January  and  February.64  But  it  seems 
more  likely  that  these  are  fish  that  either  remain 
there  throughout  the  year  or  that  visit  the  coasts 
of  New  York  and  of  southern  New  England  at 
other  times  of  year,  than  that  they  come  from 
the  Gulf  of  Maine. 

Fluctuations  in  abundance  in  the  Gulf  of  Maine. — 
Every  shore  fisherman  in  the  Massachusetts  Bay- 
Cape  Cod  region  knows  that  silver  hake  vary 
widely  in  abundance  from  year  to  year.  Catches 
by  one  set  of  six  pound-nets  at  North  Truro  on 
Cape  Cod  yielded  about  60,000  pounds  in  1946; 
237,000  pounds  in  1948;  232,000  pounds  in  1949; 
and  only  about  10,000  in  1944;  but  about  458,000 
pounds  in  1950.  Yearly  fluctuations  of  this  sort 
are  to  be  expected  at  any  given  locality,  in  the 
case  of  any  predaceous  wanderer.  And  there  is 
nothing  in  the  available  record  to  suggest  that  a 
major  alteration  has  taken  place  in  the  numbers 
of  silver  hake  in  its  center  of  abundance  in  the 
Gulf,  whether  upward  or  downward,  since  it  has 
been  an  important  fish  on  the  market. 

Occurrence  to  the  westward  and  eastward  of  the 
Gulf  of  Maine. — Silver  hake  are  described  as  abun- 
dant from  October  to  December  as  far  westward 
as  New  York,  sometimes  in  May  also,  though  few 
are  seen  there  in  summer.  And  yearly  catches  of 
some  2  to  5  million  pounds  of  "whiting"  by  pound 
nets w  suggest  that  the  beaches  of  New  Jersey 
rival  those  of  the  Cape  Cod  Bay  region  in  the 
seasonal  abundance  of  silver  hake.  But  we  have 
not  heard  of  any  great  numbers  of  them  close  in 
shore  beyond  Cape  May,  though  pound  nets  do 
take  a  few  as  far  south  as  the  mouth  of  Chesapeake 
Bay.  Farther  out  on  the  shelf,  silver  hake  of  all 
sizes  are  to  be  found  at  all  times  of  the  year,  from 
the  offing  of  southern  New  England,  westward  and 
southward,  in  numbers  large  enough  for  otter 
trawlers  to  land  3  to  5  million  pounds  yearly  in 
New  York  and  New  Jersey,56  and  smaller  amounts 
in  Delaware. 


Eastward  from  our  limits  we  find  the  silver 
hake  described  as  abundant67  in  outer  Nova 
Scotian  waters  generally.  But  we  have  no  clue  as 
to  their  actual  numbers  there,  relative  to  the  Gulf 
of  Maine,  for  they  are  not  yet  important  enough 
commercially  to  be  included  in  the  Canadian 
fisheries  statistics.  The  experimental  cruises  of 
the  Newfoundland  Fisheries  Research  Commission 
took  them  on  Banquereau  and  Misaine  Banks;  in 
the  northern  side  of  Cabot  Straits;  on  the  southern 
part  of  the  Grand  Banks ;  and  at  Bay  Bulls  on  the 
east  coast  of  the  Avalon  Peninsula,  which  is  the 
most  northern  record  for  them  of  which  we  chance 
to  know.  But  it  seems  they  are  not  known 
anywhere  in  the  Gulf  of  St.  Lawrence.68 

Importance. — Silver  hake  are  as  sweet  a  fish  as 
one  could  ask,  if  eaten  fresh  or  if  slack  salted  over 
night  and  used  for  breakfast  next  morning.  But 
they  soften  so  fast  that  there  was  no  regular 
market  demand  for  them  of  old,  and  most  of  those 
that  were  caught  incidentally  were  thrown  over- 
board. In  fact,  we  can  remember  seeing  them 
used  locally  for  fertilizer.  Thus  only  some  37,000 
pounds  were  saved  in  Maine  and  Massachusetts 
combined,  even  as  recently  as  1895.  But  improved 
methods  of  freezing  fish  were  followed  by  landings 
of  about  two  million  pounds  by  1902;  of  between 
four  and  five  million  pounds  in  1905,  rising  through 
the  years  of  the  first  world  war  to  more  than  14 
million  pounds  in  1919.69  The  yearly  landings 
then  fell  off,  for  some  reason,  to  only  about  6  million 
to  9  million  pounds  for  the  period  1924  to  1933, 
which  was  far  less  than  the  potential  catch.  But 
the  landings  then  increased  again,  as  frozen  whit- 
ing became  more  popular  in  the  Middle  West,  to 
about  15  million  pounds  in  1935,  to  about  40  mil- 
lion pounds  by  1940,  with  from  46  million  to  74 
million  pounds  during  the  6-year  period  1942  to 
1947.60 

All  but  a  small  part  of  the  Maine  and  Massa- 
chusetts landings,  recorded  in  the  following  table, 
are  from  within  the  limits  of  the  Gulf  of  Maine. 

The  silver  hake  now  ranks  fourth  or  fifth  among 
Gulf  of  Maine  fishes  in  amount  landed.     But  it 


'<  Albatross  II  trawled  8  fish,  7  to  9  Inches  long,  off  New  York,  February 
28,  1929,  at  28  fathoms;  and  the  dragger  Euaene  H.,  Capt.  Henry  Kllmm, 
picked  up  115  of  market  size  in  a  week's  trip,  about  80  miles  off  Martha's 
Vineyard,  at  47  to  67  fathoms,  January-February  1950. 

"  1942,  5,343,300  pounds;  1945,  5,842,900  pounds;  1947, 1,784,500  pounds. 

"  Otter  trawl  landings  of  "whiting,"  for  New  York  and  New  Jersey  com- 
bined, were  3,468,200  pounds  In  1942;  5,243,700  pounds  In  1945;  and  7,498,600 
pounds  In  1947.    Delaware  trawlers  reported  203,500  pounds  for  1947. 


»'  Vladykov  and  McKenzle,  Proc.  Nova  Scotia  Inst.  Scl.,  vol.  19, 1935,  p.  72. 

w  According  to  Dr.  Huntsman  all  ostensible  reports  of  their  presence  In 
the  Gulf  of  St.  Lawrence  have  been  based  In  reality  on  the  other  hakes  of  the 
genus  Urophycis  (p.  221).  And  it  is  these  that  are  meant  when  "hake"  are 
mentioned  In  the  early  publications  of  the  TJ.  S.  Fish  Commission,  such  as 
Balrd's  (Eept.  TJ.  S.  Coram.  Fish.  (1886)  1889,  app.  A.)  report  on  the  fisheries 
of  eastern  North  America. 

»  Landings  for  Maine  and  Massachusetts  combined. 

"  Maine  and  Massachusetts  combined. 


182 


FISHERY   BULLETIN   OF  THE    FISH   AND  WILDLIFE   SERVICE 


ranks  only  about  seventh  in  value.  In  1945,  the 
year  when  the  catch  was  largest,  its  value  was 
$1,736,200.  Its  rank  is  low  as  a  sportman's  fish, 
for  while  it  bites  greedily,  it  puts  up  only  a  feeble 
resistance  when  hooked. 


Year 

Catch  to 

nearest  1,000 

pounds 

Year 

Catch  to 

nearest  1,000 

pounds 

1919                 

14, 607, 000 
6,377,000 
7, 875,  000 
7, 943, 000 
6,936,000 
6, 379, 000 
8,  678, 000 
15, 420,  000 
21,038,000 

1938 

24,851,000 

1939 

27,  539, 000 

1929               

1940 _ 

39,  990, 000 

1930          

1942-. 

45,900,000 

1931           

1943 

48,460,000 

1932          .- 

1944... 

47, 373, 000 

1933           ._ 

1945 

73, 866, 000 

1935                 

1946 

48,844,000 

1937                    

1947. 

58, 936,  000 

Cod  Gadus  ccdlarias  Linnaeus  1758  6I 
.  r  ,  iwc>r)ii*£t  Rock  cod 

Jordan  and  Evermann,  1896-1900,  p.  2541. 

Description. — The  most  noticeable  external  char- 
acteristics of  the  cod,  emphasized  above  in  the  gen- 
eral survey  of  the  cod  family  (p.  173),  are  its  three 
dorsal  fins  and  two  anal  fins;  its  lack  of  fin  spines; 
the  location  of  its  ventral  fins  forward  of  its 
pectoral  fins,  and  the  fact  that  its  upper  jaw  pro- 
trudes beyond  the  lower;  that  its  tail  is  usually 
nearly  square,  and  that  its  lateral  line  is  pale, 
not  black. 

The  cod  is  a  heavy-bodied  fish,  only  slightly 
flattened  sidewise,  its  body  deepest  under  the 
first  dorsal  fin  (cod  neither  very  fat  nor  very  lean 


are  about  one-fourth  to  one-fifth  as  deep  as  they 
are  long) ,  tapering  to  a  moderately  slender  caudal 
peduncle,  and  with  a  head  so  large  that  it  takes 
up  about  one-fourth  of  the  total  length  of  the  fish. 
The  nose  is  conical  and  blunt  at  the  tip ;  the  mouth 
wide,  with  the  angle  of  the  jaw  reaching  back  as 
far  as  the  anterior  part  of  the  eye;  and  there  are 
many  very  small  teeth  in  both  jaws.  The  first 
dorsal  fin  usually  (if  not  always)  originates  well 
in  front  of  the  midlength  of  the  pectoral  fins;  it 
is  the  highest  of  the  three  dorsals,  triangular,  with 
rounded  apex  and  convex  margin.  The  second 
dorsal  fin  is  nearly  twice  as  long  as  the  first  dorsal 
and  about  twice  as  long  as  it  is  high,  decreasing 
in  height  from  front  to  rear  with  slightly  convex 
margin.  The  third  dorsal  fin  is  a  little  longer 
than  the  first  dorsal,  and  is  similar  to  the  second 
dorsal  in  shape. 

The  caudal  fin  is  about  as  broad  as  the  third 
dorsal  fin  is  long  (rather  small  for  the  size  of  the 
fish)  and  broom-shaped.  The  two  anal  fins  stand 
below  the  second  and  third  dorsals,  to  which  they 
correspond  in  height,  in  length,  and  in  shape. 
The  number  of  fin  rays  was  as  follows,  in  a  large 
series  of  Gulf  of  Maine  cod,  23  to  37  inches  long, 
examined  by  Welsh. 


•'  Jordan,  Evermann,  and  Clark  (Rept.  TJ.  S.  Comm.  Fish.  [1928],  Pt.  2, 
1930,  p.  210)  use  the  species  name  morrhua  Linnaeus  1758.  But  the  use  of 
caUarias  accords  better  with  modern  practice,  because  It  preceded  morrhua 
on  the  same  page  of  the  Systema  Naturae. 


Number  of  flnrays 

Dorsal 

Anal 

First 

Second 

Third 

First 

Second 

13 
15 
16 

19 
21 
24 

18 
19 
21 

20 
22 
24 

17 

18 

22 

Figttee  86. — Cod  (Gadus  callarias),  Eastport,  Maine.     From  Goode.     Drawing  by  H.  L.  Todd. 


FISHES   OF  THE  GULF  OF   MAINE 


183 


As  few  as  12  rays  have  occasionally  been  re- 
corded for  the  first  dorsal,  16  for  the  second,  17 
for  the  third,  17  for  the  first  anal  and  16  for  the 
second.  The  pectoral  fins,  set  high  up  on  the 
sides,  reach  back  as  far  as  the  rear  end  of  the  first 
dorsal.  The  ventral  fins  are  nearly  as  long  as  the 
pectorals  in  young  cod  but  are  shorter,  relatively 
in  large  fish,  with  the  second  ray  extending  beyond 
the  general  outline  as  a  filament  for  a  distance 
almost  one-fourth  as  long  as  the  entire  fin.  Both 
the  head  and  the  body  are  clothed  with  small 
scales. 

Young  cod  are  easily  distinguished  from  large 
tomcod  by  their  relatively  broad  ventral  fins  with 
slender  filaments,  by  the  location  of  the  first  dorsal 
fin,  and  by  their  larger  eyes,  as  explained  in  the 
description  of  that  species  (p.  196).  The  pale  lat- 
eral line  readily  distinguishes  the  cod  from  the 
haddock;  and  the  square-tipped  tail,  projecting 
upper  jaw,  and  spotted  color  pattern  of  a  cod 
give  it  an  aspect  quite  different  from  that  of  the 
pollock. 

Color. — Cod  vary  so  widely  in  color  that  sundry 
of  its  color  phases  have  been  named,  but  all  of 
them  fall  into  two  main  groups,  the  gray-green 
and  the  red.  The  back  and  upper  sides  of  the 
former  range  from  almost  black  through  dark 
sooty  or  brownish  gray,  olive  gray,  olive  brown, 
sepia  brown,  mouse  gray,  ashy  gray,  clay  colored, 
and  greenish  to  pale  pearly  (darker  on  the  back 
than  on  the  sides) ;  the  fins  are  of  the  general  body 
tint,  and  the  belly  is  whitish,  usually  tinged  with 
the  general  ground  color.  The  red  or  "rock"  cod 
vary  from  dull  reddish  brown  to  orange  or  brick 
red,  with  white  belly  tinged  with  reddish,  and 
with  red,  olive,  or  gray  fins.  In  most  cod  the 
upper  surface  of  the  body,  the  sides  of  the  head, 
and  the  fins  and  tail  (but  not  the  nose  or  belly) 
are  thickly  speckled  with  small,  round,  vague- 
edged  spots.  On  the  "gray"  fish  these  are  of  a 
brownish  or  yellowish  cast,  darker  than  the  gen- 
eral body  color,  while  they  are  usually  reddish 
brown  or  sometimes  yellowish  on  the  "red"  fish. 
Occasionally  one  sees  a  spotless  cod,  but  these  are 
unusual.  The  lateral  line  is  invariably  paler  than 
the  general  body  tint,  pearly  gray  or  reddish  ac- 
cording to  the  hue  of  the  particular  fish  in  ques- 
tion, and  it  stands  out  against  the  darker  sides. 

Size. — Cod  sometimes  grow  to  a  tremendous 
size.  A  huge  one  of  21 1  %  pounds  and  more  than  6 
feet  long,  was  caught  on  a  long  line  off  the  Massa- 


chusetts coast  in  May  1895; 62  one  that  weighed 
138  pounds  dressed  (hence  must  have  weighed  180 
pounds  or  more  alive)  was  brought  in  from  Georges 
Bank  in  1838;  and  Goode M  mentions  several 
others  of  100  to  160  pounds  as  caught  off  Massa- 
chusetts. But  cod  of  a  hundred  pounds  are 
exceptional,  the  largest  New  England  cod  of 
which  we  have  heard  recently  being  one  of  90 
pounds,  that  was  taken  off  the  coast  of  Maine 
early  in  July  1922.  Even  a  75-pound  fish  is  a 
rarity,  but  50  to  60  pounders  are  not  unusual. 
The  so-called  "large"  fish  that  are  caught  near 
shore  run  about  35  pounds;  and  "large"  ones  taken 
on  Georges  Bank  about  25  pounds.  But  the  shore 
fish,  large  and  small  together,  average  only  be- 
tween 6  and  12  pounds  in  weight. 

The  relationship  between  length  and  weight  is 
usually  about  as  follows  for  fish  caught  on  the  in- 
shore grounds  between  Cape  Ann  and  Portland, 
though  this  varies  with  the  condition  of  the  fish 
and  with  their  state  of  sexual  development.64 


Females 

Males 

Inches 

Pounds 

Inches 

Pounds 

19  to  20- 

2W-3 

3W-  4 

4H-  7 

5-7 

7-9 

7H-10 

9    -13 

12H-17H 

16    -23 

18    -22 

16    -32 

29H-32 

31     -51 

50 

54 

20  to  21 

3    -3H 
4-5)4 
6M-  8 
7    -8M 

23  to  24 

23  to  24 

25  to  26 

25  to  26. 

27  to  28 

27  to  28 

30  to  31 

30  to  31.. 

32  to  33 

32  to  33.  _ 

34  to  35 

34  to  35 

36  to  37 

1254-17 

36to36H 

38  to  39 

38  to  39 

40  to  41 

40  to  42 

43  to  45 

25VS-29 

43  to  44 

46 

48^  to50H 

52 

57^ 

A  99%-pound  fish  recorded  by  Earll  was  62 
inches  long,  and  one  of  100  pounds  caught  off 
Wood  Island,  Maine,  on  April  9,  1883,  measured 
65  inches,  its  head  17%  inches.  Any  fish  of  5%  to  6 
feet  will  weigh  100  pounds  or  more. 

Habits. — Cod  in  one  place  or  another  range  from 
the  surface  down  to  250  fathoms  at  least. 

During  the  first  year  after  the  young  cod  take  to 
bottom  (p.  186)  many  of  them  five  in  very  shoal 
water,  even  along  the  littoral  zone,  and  many 
young  fry  have  been  taken  at  Gloucester  and  else- 
where along  the  shores  of  New  England,  while 

u  Jordan  and  Evermann,  American  Food  and  Game  Fishes.     1902,  p.  514. 

•»  Fish  Ind.  U.  8..  Sec.  1,1884,  p.  220. 

M  Based  chiefly  on  measurements  given  by  Earll  (Kept.  XJ.  3.  Comm.  Fish. 
[1878|,  1880,  p.  734),  and  on  a  large  series  of  cod  measured  fresh  from  the  nets  by 
Welsh  during  the  spring  of  1913. 


184 


FISHERY  BULLETIN   OF   THE    FISH   AND  WILDLIFE   SERVICE 


many  small  cod  are  caught  about  the  rocks  only  a 
fathom  or  two  deep  even  in  summer.  But  it  is 
certain  that  many  cod  fry  take  to  bottom  on  the 
offshore  banks  also,  for  we  have  trawled  young  fry 
at  many  localities  between  Nantucket  Shoals  and 
Browns  Bank.  As  a  rule,  the  large  cod  lie  deeper 
than  7  or  8  fathoms  in  summer  in  our  latitudes. 
But  the  fishing  is  often  good  in  only  3  to  5  fathoms 
of  water  in  wintertime,  especially  in  Ipswich 
Bay.  At  the  other  extreme,  comparatively  few 
cod  are  caught  much  deeper  than  100  fathoms 
within  the  Gulf  of  Maine.  And  although  fisher- 
men sometimes  do  well  at  much  greater  depths  on 
the  slopes  of  the  offshore  banks,  the  5-  and  75- 
fathom  contours  probably  include  the  great 
majority  of  all  the  cod  living  in  the  Gulf,  summer 
or  winter. 

The  largest  catches  of  cod  are  made  on  rocky 
and  pebbly  grounds;  on  gravel;  on  sand,  and  on 
a  particularly  gritty  type  of  clay  with  broken 
shells.  They  also  frequent  the  deeper  slopes  of 
ledges  along  shore,  where  they  forage  among  the 
Irish  moss  (Chondrus  crispus)  and  among  sea- 
weeds of  other  kinds.  Young  red  ones  are  espe- 
cially common  in  these  situations,  while  one  some- 
times catches  a  large  rock  cod  as  these  dark  brown 
or  red  fish  are  called.  And  the  bottoms  where 
cod  and  hake  are  found  are  so  distinct  that  a  long 
line  set  from  a  hard  patch  out  over  the  soft  sur- 
rounding ground  will  often  catch  cod  at  the  one 
end,  hake  at  the  other.  But  fair  catches  are 
sometimes  taken  on  mud,  as  off  Mount  Desert, 
where  large-  and  medium-sized  cod  are  regularly 
caught  on  soft  ground  in  winter.  And  a  few  very 
large  cod  (35-60  lb.)  have  also  been  brought  in 
from  the  mud  bottom  of  the  deep  basin  to  the 
westward  of  Jeffreys  Ledge  (about  90  fathoms). 

The  cod,  as  appears  from  the  foregoing,  is  typi- 
cally a  ground  fish;  except  on  some  journey  (a 
subject  to  be  discussed  later)  or  when  following 
its  prey,  it  usually  lies  within  a  fathom  or  so  of  the 
bottom.  And  large  ones  keep  closer  to  the  ground 
than  small  ones  as  a  rule,  so  that  the  closer  one 
fishes  to  bottom  the  larger  the  cod  are  likely  to 
run.  But  even  the  large  ones  sometimes  follow 
herring  up  to  the  surface;  we  have  known  of 
large  cod  gaffed  from  a  vessel's  side  in  Northeast 
Harbor,  Mount  Desert  Island,  in  September, 
while  they  were  chasing  sardines.  And  they  come 
to  the  surface  more  commonly  on  the  Grand 
Banks  and  along  the  eastern  coast  of  Labrador, 


when  they  are  following  capelin.  Cod  even 
strand  on  the  Labrador  beaches  while  harrying 
schools  of  capelin,  but  we  have  never  known  cod 
to  strand  anywhere  around  the  coasts  of  the 
Gulf  of  Maine,  as  silver  hake  so  often  do  (p.  175). 

The  adult  cod  is  at  home  in  any  temperature 
from  32°  to  50°-55°  F.;  in  all  but  the  superficial 
layers  of  the  Gulf  of  Maine,  that  is,  at  all  seasons. 
But  experience  at  the  Woods  Hole  hatchery, 
proves  that  freezing  may  be  fatal  by  the  forma- 
tion of  anchor  ice.  On  the  other  hand,  while 
large  cod  tend  to  avoid  water  warmer  than  about 
50°  F.,  except  that  they  are  abundant  at  times 
in  temperatures  as  high  as  58°-59°  F.  on  Nan- 
tucket Shoals  (the  most  southerly  year-round 
cod-ground  in  the  Atlantic).  Small  cod  are 
somewhat  less  sensitive  to  heat  than  large,  a  fact 
reflected  in  the  presence  of  greater  numbers  of 
them  in  shoal  water  in  summer  than  of  larger 
fish.  The  relationship  of  the  spawning  of  the  cod 
to  temperature  is  discussed  below  (p.  194). 

Food.— When  the  larval  cod  first  breaks  from 
the  egg  it  subsists  on  the  yolk  with  which  its 
abdomen  is  distended  (fig.  8S),  as  do  most  other 
sea  fishes.  But  this  source  of  nutriment  is  com- 
pletely absorbed  by  the  sixth  day  after  hatching, 
and  the  future  existence  of  the  little  fish  depends 
as  much  on  finding  a  plentiful  supply  of  food  as 
on  escaping  the  enemies  by  which  it  is  encom- 
passed. So  far  as  known,  the  larval  and  post- 
larval  cod  subsist  almost  exclusively  on  copepods 
and  on  other  minute  Crustacea,  during  the  several 
months  while  they  are  drifting  in  the  upper  layers 
of  water.96  And  this  same  diet,  varied  with 
amphipods,  barnacle  larvae,  and  other  small 
crustaceans,  as  well  as  with  small  worms,  is  the 
chief  dependence  of  the  little  cod  when  they  first 
seek  the  bottom  M  but  as  they  grow  larger  they 
consume  invertebrates  in  great  variety  and  in 
enormous  amount. 

Mollusks,  collectively,  are  probably  the  largest 
item  in  the  cod's  diet  in  the  Gulf  of  Maine;  any 
shellfish  that  a  cod  encounters  is  gobbled  up,  so 
that  their  stomachs  are  mines  of  information  for 
students  of  mollusks.     Large  sea  clams  {Mactra), 


"  Bumpus,  Science.,  N.  Ser.,  vol.  7, 1898,  p.  485. 

"  For  further  details  on  the  diet  of  cod  larvae  and  fry,  see  Brook  (5  ann. 
Rept.,  Fish.  Board  Scotland  (1886)  1887,  p.  327),  Mcintosh  and  Masterman 
(British  Marine  food  fishes,  1897,  p.  242),  Kendall  (Rept.  U.  S.  Comm.  Fish. 
(1896)  1898,  p.  179),  Bumpus  (Science,  N.  Ser.,  vol.  7, 1898  p.  485),  and  Good- 
child,  Graham  and  Carruthers  (British  Mlnlst.  Agrlc.  Fish.,  Fish.  Inv., 
Ser.  2,  vol.  8,  No.  6,  [1925]  1926. 


FISHES   OF  THE  GULF   OF   MAINE 


185 


the  empty  shells  of  which  are  often  found  neatly 
nested  in  cod  stomachs :  cockles  (Polynices) ;  and 
sea  mussels  (Modiolus)  are  staples,  all  of  which 
they  swallow  whole.  Cod  also  eat  crabs,  hermit 
crabs,  lobsters  (large  and  small),  shrimps,  brittle 
stars  (of  which  they  are  sometimes  crammed  full) , 
sea  urchins,  sea  cucumbers,  and  sea  worms 
(Nereis).  Brittle  stars  and  small  crabs,  for 
example,  had  been  the  chief  diet  of  the  cod 
examined  by  Welsh  on  the  Isles  of  Shoals-Boon 
Island  ground  in  April  1913,  while  Wilcox  67  states 
that  a  number  of  17-pound  fish  caught  in  Ipswich 
Bay  were  full  of  large  red  prawns  2  to  4  inches 
long  (evidently  the  northern  edible  shrimp 
Pandalus).  And  we  have  found  crabs  (Cancer; 
Libinia)  the  chief  food  of  the  cod  on  Nantucket 
shoals. 

Tunicates  (sea  squirts)  also  bulk  large  in  then- 
diet.  Occasionally  they  eat  hydroids,  bryozoans, 
and  algae,  perhaps  taking  these  for  the  amphipods 
that  are  hidden  among  them.  And  in  late  summer 
cod  frequently  feed  on  ctenophores  (Pleurobrachia 
fileus).  But  while  its  diet  list  would  probably 
prove  almost  as  extensive  as  that  of  the  haddock 
(p.  202),  the  cod  shows  so  decided  a  preference  for 
large  shells  rather  than  for  small  ones  that  the 
stomach  contents  of  cod  and  haddock  taken  side 
by  side  differ  noticeably.  Nor  is  it  likely  that 
cod  root  the  bottom  as  haddock  do  (p.  202),  for 
worms. 

Cod  pursue  and  gorge  on  squid  at  every  oppor- 
tunity, and  on  various  small  fish,  particularly  on 
herring,  on  launce,  and  (in  the  north)  on  capelin; 
also  on  shad,  mackerel,  menhaden,  silversides, 
alewives,  silver  hake,  young  haddock,  and  even 
on  their  own  young,  rising  into  the  upper  waters 
for  this  purpose  when  necessary  (p.  184).  They 
also  pick  up  flounders,  cunners,  rock  eels  (Pholis) , 
blennies,  sculpins,  sea  ravens,  small  hake  and 
skates  from  the  bottom.  In  fact,  they  take  any 
fish  small  enough  to  swallow,  including  the  hard 
slim  alligatorfish  (p.  457)  and  even  the  sea  horse 
(p.  315).  And  Welsh  noted  that  many  cod  taken 
near  the  Isles  of  Shoals  on  May  1,  1913  spat  up 
small  rosefish  from  4  to  6  inches  long.  The  eggs 
of  the  longhorn  sculpin M  and  of  the  eelpout 
(Macro zoarces)  69  also   have   been   found   in   cod 

"  Bull.  U.  S.  Fish.  Comm.,  vol.  6, 1887,  p.  95. 
■  Warfel  and  Merrinmn,  Copeia,  1944,  p.  198. 

•'  Olsen  and  Merrlman,  Bull.  Bingham  Oeeanogr.  Coll.,  vol.  9,  art.  4, 
1946,  p.  77. 

210941—53 13 


stomachs.  Adult  cod  as  well  as  small  are  also 
known  to  feed  on  pelagic  shrimps  in  the  waters 
around  Iceland,70  but  we  have  never  heard  of 
them  doing  so  in  the  Gulf  of  Maine. 

Even  a  wild  duck  does  not  escape  from  a 
large  cod  now  and  then.  Thus  we  have  heard  of 
several  scoters  found  in  the  stomachs  of  large  fish 
caught  off  Muskeget  Island  in  1897;  and  though 
sea  fowl  are  not  a  normal  article  in  their  diet,  the 
flesh  of  the  greater  shearwater  (hagdon)  has  long 
been  considered  excellent  cod  bait.  Objects  as 
indigestible  as  pieces  of  wood  and  rope,  fragments 
of  clothing,  old  boots,  jewelry,  and  other  odds  and 
ends  have  repeatedly  been  found  in  cod  stomachs. 
And  they  often  swallow  stones;  but  probably  for 
the  anemones,  hydroids,  and  other  animals  growing 
thereon,  and  not  to  take  on  ballast  for  a  journey 
as  the  old  story  has  it. 

Although  cod  are  so  rapacious  they  fast  gener- 
ally while  they  are  spawning;  the  stomachs  of 
nearly  all  the  ripe  fish  examined  by  Earll,  and 
recently  by  Welsh,  were  empty. 

Experiments  performed  on  the  cod  in  captiv- 
ity,71 combined  with  the  general  experience  of 
fishermen,  suggest  that  they  capture  moving 
objects  by  sight.  But  apparently  cod  (and  for 
that  matter  other  fish),  can  see  clearly  only  for  a 
few  feet,  and  their  greediness  in  snapping  up  the 
naked  meat  of  clams  and  cockles  (foods  which 
they  never  find  in  that  condition  in  nature) ,  added 
to  the  fact  that  they  bite  as  readily  by  night  as 
by  day,  seems  sufficient  evidence  that  they 
depend  largely  on  smell. 

Enemies. — In  the  Gulf  of  Maine,  large  sharks 
and  the  spiny  dogfish  are  the  worst  enemy  of  the 
adult  cod.  Formidable  enemies  of  young  cod 
fry  are  the  small  pollock  which  infest  our  harbors. 
These  are  so  fierce  that  a  single  pollock  7  or  8 
inches  long  will  disperse  a  school  of  hundreds  of 
cod  fry,  driving  them  to  shelter  among  the  weeds 
and  rocks,  while  Earll  remarks  that  in  the  aquar- 
ium a  cod  so  fears  a  pollock  of  equal  size  that  it 
will  invariably  hide  if  possible.  Young  cod,  up  to 
7  to  8  inches,  are  also  devoured  in  large  numbers 
by  the  larger  cod. 


n  Schmidt  (Skrift.  Komm.  Havunderstfgelser,  No.  1,  1904,  p.  70)  and 
Paulsen  (Meddelel.  Kommls.  Havunderstfgelser,  Serie  Plankton,  vol.  1, 
No.  8,  1909,  p.  39). 

«  Bateson,  Jour.  Mar.  Biol.  Assoc.  United  Kingdom,  N.  Ser.,  vol.  1, 
1889-90,  p.  241. 


186 


FISHEKY    BULLETIN    OF   THE    FISH   AND    WILDLIFE    SERVICE 


Migrations  and  wanderings. — It  has  long  been 
known  that  cod  carry  out  extensive  migrations 
in  some  regions,  but  that  they  are  more  nearly 
stationary  in  others.  European  (especially  the 
Scandinavian)  biologists  have  succeeded  in  tracing 
the  major  outlines  of  their  movements  for  North 
European  seas,  and  enough  evidence  has  accumu- 
lated to  show  that  their  travels  fall  into  the  same 
categories  in  the  one  side  of  the  Atlantic  as  in 
the  other.  These  categories  are:  (a)  involuntary 
drifts  by  the  eggs  and  by  the  larvae  before  they 
take  to  the  bottom;  (b)  the  various  journeyings 
by  the  older  cod  in  search  of  food;  (c)  journeys 
associated  with  the  concentrations  of  cod  on 
particular  spawning  grounds;  and  (d)  regular 
seasonal  migrations  (with  return  movement) 
between  different  regions  that  are  suitable  for 
cod  during  different  parts  of  the  year. 

To  begin  with,  the  eggs,  larvae,  and  young  fry 
of  the  cod,  bike  those  of  so  many  other  sea  fishes, 
drift  helplessly  with  the  current  from  the  time 
they  are  spawned  until  they  seek  the  bottom  (a 
fact  established  by  European  observations  too 
numerous  to  list).72  The  length  of  this  period 
(varying  in  duration  in  different  seas)  depends 
partly  on  whether  the  fry  are  near  land  or  are 
far  out  at  sea,  and  partly  on  whether  they  are 
floating  over  deep  water  or  over  shoal.  It  is  not 
likely  to  last  for  more  than  two  months  for  fish 
that  are  hatched  on  the  inshore  spawning  grounds 
in  the  Gulf  of  Maine,  where  the  bottom  is  within 
easy  reach.  Even  so,  it  is  extremely  unlikely 
that  any  cod  fry  take  to  the  bottom  near  where 
they  were  spawned. 

This  matter  is  discussed  further  in  relation  to 
the  occurrence  of  the  cod  in  our  Gulf  (p.  190). 

The  journeyings  of  th